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THE BIOLOGICAL SOCIETY OF WASHINGTON
1999-2000
Officers
President: Richard P. Vari Secretary: Carole C. Baldwin
President-elect: Brian F. Kensley Treasurer: T. Chad Walter
Elected Council
Michael D. Carleton Rafael Lemaitre
W. Duane Hope Roy W. McDiarmid
Susan L. Jewett James N. Norris
Custodian of Publications: Storrs L. Olson
PROCEEDINGS
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EDITOR
BIOLOGICAL SOCIETY OF WASHINGTON
NATIONAL MUSEUM OF NATURAL HISTORY
WASHINGTON, D.C. 20560, U.S.A.
Known office of publication: National Museum of Natural History, Smithsonian Institution,
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This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
This number is dedicated to
Dr. Austin B. Williams
Systematic Zoologist
National Systematics Laboratory
National Marine Fisheries Service
National Museum of Natural History
Smithsonian Institution
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):1—12. 2000.
Austin Beatty Williams (17 October 1919-27 October 1999).
Biographical summary
Rafael Lemaitre and Bruce B. Collette
(RL) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560-0163, U.S.A.;
(BBC) Bruce B. Collette, National Marine Fisheries Service Systematics Laboratory, National
Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0153, U.S.A.
Abstract.—The professional career and aspects of the life of Austin Beatty
Williams (17 October 1919-27 October 1999), Systematic Zoologist for the
Systematics Laboratory of the U.S. National Marine Fisheries, are summarized.
Included is a bibliography with the 118 papers published by Williams, and a
list of all the new names he proposed along with the holotype repository and
catalogue number of species and subspecies.
Austin B. Williams (Fig. 1) was System-
atic Zoologist at the Systematics Laborato-
ry, National Marine Fisheries Service
(NMES), based at the National Museum of
Natural History, Smithsonian Institution,
Washington, D.C. After a valiant fight with
cancer, Austin passed away at his home in
Falls Church, Virginia. Shortly after his
death, the Council of the Biological Society
of Washington unanimously voted to dedi-
cate this issue, the first of the new millen-
ium, to his memory. This dedication is most
fitting as Austin generously served the So-
ciety 1n many capacities during the last
quarter of the 20th century, and gave luster
to the Proceedings by using it to publish
many of his important papers. He served
the Society as a Editor of the Proceedings
(1974-1977), Vice-president (1983-1986),
President (1986-1988), Past-President
(1989-1999), Custodian of Publications
(1989-1995), and contributed significantly
to its financial soundness as a member of
the Finance Committee (1995-1999). He
was editor of Bulletin No. 3: ““Symposium
on the Composition and Evolution of Crus-
taceans in the Cold and Temperate Waters
of the World Oceans”’ (1979), based on the
results of a U.S.-U.S.S.R. Cooperative Pro-
gram. He also provided the summary chap-
ter for Bulletin No. 6: ‘““The hydrothermal
vents of the eastern Pacific: An overview”
(Williams 1985b).
Austin had a distinguished career span-
ning five decades during which he pub-
lished 118 papers (see bibliography). Born
in Plattsburg, Missouri in 1919, he was the
first child of Oliver Perry Williams and
Lucy Sell; his siblings are brothers Hillis
and Oliver. He married Jean McNicol with
whom he had their only child, David (mar-
ried to Anita Kyle, with two children, Lau-
ren and Kyle). His family had only modest
means so he had to work to support his ed-
ucation, first at McPherson College (A.B.
1943), and then at the University of Kansas
(Ph.D. 1951), where he studied Ozark cray-
fishes. His studies on these crayfishes re-
main among the key references to identify
these decapods in the region. From 1951 to
1955 he was with the University of North
Carolina Institute of Fisheries Research,
studying the life history and ecology of
penaeid shrimps. He then worked at the
University of Illinois from 1956 to 1963,
after which he returned to the North Caro-
lina Institute of Fisheries Research to con-
tinue his studies on marine and estuarine
decapods. In the mid 1960s, Donald F
Squires, Chairman of the Department of In-
i)
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
at Heart Tail Ranch, Butte County, South Dakota, one of his favorite fossil collecting sites, 31 July 1997, in
back is son David, in front left to right are David’s wife Anita with grandchildren Kyle and Lauren, and Norma
Samuels (Norma Samuels); recording observations in his notebook on fossil Cretaceous decapods collected from
the Pierre Shale at Heart Tail Ranch, Butte County, South Dakota, 6 August 1997 (Gale A. Bishop); at Mount
Rushmore, South Dakota, August 1997 (Norma Samuels).
vertebrate Zoology and then Deputy Direc-
tor of the National Museum of Natural His-
tory, Smithsonian Institution, considered
him for a job; however, his interview with
Secretary Dillon Ripley did not go well
(most probably because of Austin’s non-as-
suming personality), and he turned him
Austin B. Williams. Clockwise from upper left: from church directory, 1995; with family and friends
down. This was a loss for the Museum but
fortunately he was hired by the NMFS’
Systematics Laboratory in 1971.
In addition to his crustacean work, Aus-
tin served NMFS and the Museum in a
number of different ways. For example, he
represented the Allied Agencies (NMFS,
VOLUME 113, NUMBER 1
Agriculture, and what was then the Fish and
Wildlife Service) on the Senate of Scientists
in the Museum. During his tenure in this
position, a question arose as to whether or
not the administrative staff of the Museum
had increased significantly in the several
preceding years. Discussion went back and
forth between the Senate and the Director
of the Museum. Finally, Austin volunteered
to get some real data on the issue. In a typ-
ical A. B. Williams way, he systematically
went through the entire telephone directory
and counted museum administrators at 5-
year intervals, and demonstrated that the
type of positions that the Senate considered
as ‘“‘administrative’’ had in fact increased
significantly.
He was the acknowledged expert on and
leader in studies of the systematics of east-
ern American decapod crustaceans. He is
probably best known for his widely used
monograph “Shrimps, lobsters, and crabs
of the Atlantic coast of the eastern United
States’’ published by the Smithsonian in
1984. His earlier study on the decapods of
the Carolinas published in 1965, a precursor
to his 1984 monograph, was selected as a
Science Citation Classic in 1983, a rare
honor for a systematist. His invaluable pa-
per (Williams 1987a) on the identification
of spiny lobsters by color patterns of the
tails grew into a book co-authored with I.
Dore, entitled ‘‘Lobsters of the world—an
illustrated guide’? (Williams & Dore
1988e); these two publications are indis-
pensable for anybody interested in this
group of economically significant decapods.
The primary focus of his research was
the taxonomy, systematics, biogeography
and evolution of various decapod groups,
both fossil and Recent. He named 101 new
decapod taxa (see list), including one su-
perfamily, 2 families, 16 genera, 80 species,
and 2 subspecies. Occasionally he also
worked on other groups such as cirripeds,
mysids, amphipods, and euphausiids, and
even bird ecology. His publications provide
us with a standard of excellence, and are
well known for attention to detail, accuracy,
and usefulness in the identification of spec-
imens while at the same time giving insight
into phylogenetic relationships. No major
group of decapods escaped Austin’s atten
tion. He published important works on
crayfishes, peneaeoids, carideans, thalassin-
ideans, lobsters, anomurans, and brachyu-
rans. His landmark studies on swimming
crabs of the genus Callinectes, mud shrimps
of the family Upogebiidae, commercial lob-
sters, xanthid crabs, and deep-sea hydro-
thermal vent decapods, among others, have
earned him a place in the history of Zool-
ogy. His contributions to the systematics of
hydrothermal vent decapods inspired other
colleagues, and one genus and species of
the crab family Bythograeidae Williams,
1980, was named after him (Austinograea
williamsi Hessler & Martin, 1989, Journal
of Crustacean Biology 9(4):645—-661). In
March of 1996 he traveled to Kumamoto,
Japan, on a Japanese fellowship program to
join Keiji Baba (Kumamoto University Fac-
ulty of Ecucation) in the study of galatheids
and other vent decapods from hydrother-
mally active sites in the western Pacific. Al-
though his work concentrated on aspects of
systematics and evolution, he also pub-
lished key studies on the biology of com-
mercial penaeid shrimps, ecology of mero-
plankton, larval genetics, and crustacean
fisheries and mariculture. At the time of his
death he had completed work but unfortu-
nately left unpublished, an important revi-
sion of the crab family Latreilliidae.
Austin’s impact on carcinology is not
limited to that derived from his publica-
tions. Throughout his exemplary career he
actively participated in many professional
societies in addition to the Biological So-
ciety of Washington. He was president of
the Atlantic Estuarine Research Society
(1960-1961); co-founder, secretary (1971-—
1973), and president (1983-1985) of the
Estuarine Research Federation; secretary
(1985-1988) for the Society of Systematic
Biology; associate editor for The Crusta-
cean Society (1986-1991); and vice-presi-
dent (1990-1991) and president (1991-—
+ PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
1992) of the American Association for Zoo-
logical Nomenclature. He was also a mem-
ber of the American Association for the
Advancement of Science, American Fish-
eries Society, American Institute of Biolog-
ical Sciences, American Institute of Fishery
Research Biologists, American Society of
Limnology and Oceanography, Society for
Integrative and Comparative Biology (for-
merly American Society of Zoologists), As-
sociation of Systematic Collections, Ecolog-
ical Society of America, Kansas Academy
of Science, and Society for the Study of Evo-
lution. His editorial activities with various
journals and symposia proceedings pro-
duced many significant contributions, and
his involvement with doctoral and master-
level students at various academic institu-
tions led to the development of outstanding
carcinologists who now must continue his
legacy. His skills as reviewer were highly
regarded by editors and grant program man-
agers who were assured of a detailed, un-
biased evaluation.
Several of Austin’s papers won important
awards or honorable mentions. His lobster
identification paper (Williams 1987a), for
example, won the highly regarded ‘*Nation-
al Marine Fisheries Service Outstanding
Publication Award” for best paper in the
1997 Marine Fisheries Review. In recogni-
tion of Austin’s life-time work, The Crus-
tacean Society presented him in 1997 with
their ““Excellence in Research Award”’ dur-
ing a ceremony at the National Museum of
Natural History, Smithsonian Institution,
Washington, D.C. [see Lemaitre, R., 1998,
Journal of Crustacean Biology 18(3):619-—
620]. He was also honored with the ‘‘1999
Elton Sette Award” from the Marine Fish-
eries Section of the American Fisheries So-
ciety.
Austin will be remembered not only for
his impressive scientific accomplishments
but also for his human qualities. Unselfish
almost to a fault, he made every effort to
help colleagues and students alike. His
height of 1.85 m (6'1”) gave him a towering
physique which combined with his deep
knowledge of decapods made him an im-
posing figure to both students and junior
colleagues; however, his modesty was such
that he made sure to treat everyone as a
friend or colleague of similar stature. He
accepted life-time honors bestowed upon
him by his peers only hesitantly, and main-
tained until his end that he was undeserving
of such attention. His personality, working
habits, discipline, and inspirations date back
to his early life experiences which he often
mentioned to friends during casual conver-
sations. One of his first jobs prior to enter-
ing college was at his family’s farm in Ster-
ling, Colorado, where he helped string fenc-
es, some of which had to be modified to
‘first class communications grade’”’ by in-
sulating the top strand of barbed wire to
carry telephone signals. This worked fine
until it rained and the circuits became
grounded by water. During his Sterling days
he also taught high school. One of Austin’s
scientific strengths was his observational
and note taking abilities. He had been
trained at Kansas to write reflective notes
each night as if they were to be published.
Those who have examined his field and of-
fice notebooks are struck by how remark-
ably clear and detailed they are.
Austin often mentioned the impact of the
Great Plains of Kansas and Colorado on his
psyche. One of his closest friends, Gale A.
Bishop (Georgia Southern University), has
said that he was impressed with Austin’s
collegiality when he first met him during a
visit to the Smithsonian to study fossil
decapods. Gale suggested that he might
want to join him in the field in South Da-
kota to collect fossils. Austin did so with
much enthusiasm, and the two worked to-
gether almost every summer from 1980 un-
til the year of his death. Nancy Brannen
Marsh (Science Department, Portal High
School, Georgia) also joined them, and the
three collaborated in studies of decapods of
the Western Interior Cretaceous, collecting
numerous fossil crabs, lobsters, and shrimps
from the Carlile Shale and the Pierre Shale
of South Dakota, Wyoming and Colorado.
VOLUME 113, NUMBER 1
These fossil collections have been donated
to the Museum of Geology at the South Da-
kota School of Mines, Rapid City. He con-
fessed to Gale that the timing of their col-
laboration was most appropriate as his wife
Jean had passed away (1983) after a diffi-
cult illness (with Austin as major care giv-
er), and his return to the Great Plains was
just what he needed to gain closure and
healing from the loss. This theme, Gale
says, ““came up many times and we con-
cluded that getting back to our roots was an
extremely healing process, both for Austin’s
loss and for the loss of both of my parents;
it was an annual ‘rehealing’ as we came
back into harmony with our roots and
Mother Earth. When working with Austin
on fossils in the Western Interior his intel-
lect and collegiality were always apparent.
His interests spanned the sciences, arts, ed-
ucation, and humanities. He often would
visit the Rapid City Astronomy club to par-
ticipate in telescopic observations, take us
all out dancing at the Broken Boot Saloon
in Rapid City, climb Bear Butte or Harney
Peak, or head us up to Rushmore for the
evening patriotic lighting program. While
in the field, he brought new insights to pa-
leontology, often seeing things we took for
granted or forcing clearer explanation of
our mutual deductions. Our collaboration
was clearly very beneficial to Austin as
well as to paleontologists Nancy Brennan
Marsh, the late Reinhard Forster of Munich,
and Georgia Southern students Mike Klug,
Mehmet Samiratedu, and Amy Samiratedu.
These insights were carried over into the
laboratory and into collaborations on papers
and research comparing Recent and fossil
decapods.”’
The multi-faceted personality of Austin
included a deep appreciation of the simple
things of life, his family and friends. During
the last decade or so of his life he was for-
tunate to share many moments with Norma
Samuels, of Fairfax, Virginia, whose com-
panionship undoubtedly enriched his life.
Austin developed a passion for ballroom
and international dancing, and a love for
choral music. He actively participated in
several choral groups, including the Wash-
ington Cathedral Choral Society which per-
formed in the National Cathedral, Washing-
ton, D.C. One of his performances is pre-
served on an audio CD-ROM entitled ‘‘Mil-
lenium; Russian Choral Music’’ (1990
Centaur Record Inc.). He worshipped, sang
and was an active member of the New York
Avenue Presbyterian Church, of Washing-
ton, D.C., where well-attended and emo-
tional services were held for him on Octo-
ber 30, officiated by The Rev. Robert H.
Craig.
Austin’s remains are buried in Marion,
Kansas, alongside those of his wife.
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VOLUME 113, NUMBER 1
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8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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waters influenced by hydrothermal discharge,
brine, and hydrocarbon seepage.—Fishery Bul-
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1988d.
1988e.
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, & P. J. B. Scott. Upogebia corallifora,
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» Le Ge Abele;-D. L. Feldes eat
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, & J. J. McDermott. An eastern United
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, & K. Baba. New squat lobsters (Gal-
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Arc Basin, East Pacific Rise, and Cascadia Ba-
sin.—Fishery Bulletin 87(4):899-910.
1990c. Kensley, B., & . Axlopsis eximia, a new
thalassinidian shrimp (Crustacea, Decapoda,
Axiidae) from the Middle Eocene of South Car-
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, & E. E. Boschi. Panopeus margentus,
a new crab from the Argentine warm temperate
subregion (Decapoda: Xanthidae).—Proceed-
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103(3):598—-601.
, & N. Ngoc-Ho. Pomatogebia, a new
genus of thalassinidean shrimps from western
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Proceedings of the Biological Society of Wash-
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, & R. W. Heard. Upogebia spinistipula,
a new burrowing shrimp from the Florida shelf,
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1989b.
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VOLUME 113, NUMBER 1
gebiidae).—Proceedings of the Biological So-
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, & R. B. Moffitt. Crabs from the Mar-
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1991c. Bishop, G. A., &
sonorum, new species, a crab (Decapoda: Ca-
lappidae) from the Cretaceous Carlile Shale
(Turonian), Western Interior United States.—
Journal of Crustacean Biology 11(3):451—459.
. Comments on the proposed conser-
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cana Kellogg, 1906 (Crustacea, Branchiopoda).
(Case 2728; see BZN 47:178—-183; 48:57).—
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1991le. Van Dover, C. L., & . Egg size in squat
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, & R. Wahle. Distinguishing juvenile
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. Review: Decapod Crustacea of the At-
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. Review: Marine Lobsters of the World.
An Annotated and Illustrated Catalog of Species
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Synopsis No. 125, vol. 13:292 pp. Rome.—The
Lobster Newsletter 5:14.
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pensis Krauss, 1843 and G. africana Ortmann,
1894 (currently Upogebia capensis and U. af-
ricana; Crustacea, Decapoda) by the designa-
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(Case 2827; BZN 49(3):187—190.).—Bulletin
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. Mud shrimps, Upogebiidae, from the
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199 1d.
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L9D2c.
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1993b.
1593¢.
1994.
(Brachyura: Majidae).—Crustacean Research
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1995a. Rodriguez, G., & . Epilobocera weth-
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capoda: Brachyura: Pseudothelphusidae) from
Hispaniola.—Proceedings of the Biological So-
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1995b. Kornfield, [., , & R. S. Steneck. As-
signment of Homarus capensis (Herbst, 1792),
the Cape lobster of South Africa, to the new
genus Homarinus (Decapoda: Nephropidae).—
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, & EC. Dobbs. A new genus and spe-
cies of caridean shrimp (Crustacea: Decapoda:
Bresiliidae) from hydrothermal vents on Loihi
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1995e. Pequegnat, L. H., & . Two new species
of Munidopsis (Decapoda: Anomura: Galathei-
dae) from the Western Atlantic Ocean.—Journal
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. Comments on the proposed conser-
vation of the generic names Monstrilla Dana,
1849 and Thaumaleus Kréyer, 1849 (Crustacea,
Copepoda) (Case 2894; see BZN 52(3):245-—
249).—Bulletin of Zoological Nomenclature
53(2):122-123.
. Comments on the proposed conser-
vation of the generic name Glomeris Latreille,
1802 (Diplopoda) and the specific name of Ar-
madillo vulgaris Latreille, 1804 (Crustacea, Is-
opoda), and the application for a ruling on the
status of the name Armadillo Latreille, 1802
(Crustacea, Isopoda). (Case 2909; see BZN
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menclature 53(2):121—122. [Resolved in Opin-
ion 1897, Bulletin of Zoological Nomenclature
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Bythograea Williams,
1980. Arthropoda, Crustacea, Decapoda, Brach-
yura, Brachyrhyncha, Bythograeoidea, Bytho-
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Bahamian Field Station, June 9—13, 1995.
. Occurrence of three species of mud
shrimps in aquaculture ponds on Caribbean
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1995¢:
1995d.
1996a.
1996b.
1997a.
thermydron
1997c.
10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
description of Upogebia omissago Williams,
1993 (Decapoda: Upogebiidae).—Proceedings
of the Biological Society of Washington 110(3):
412-416.
. Two new species and a range exten-
sion of mud shrimps, Upogebia, from Pacific
Costa Rica and Mexico (Decapoda: Thalassin-
idea: Upogebiidae).—Proceedings of the Bio-
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1998a. Baba, K., & . New Galatheoidea (Crus-
tacea, Decapoda, Anomura) from hydrothermal
systems in the West Pacific Ocean: Bismarck
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ma 20(2):143—156.
, & J. L. Hernandez-Aguilera. A new
species of mud shrimp, Upogebia toralae, from
Veracruz, Mexico (Decapoda: Thalassinidea:
Upogebiidae).—Proceedings of the Biological
Society of Washington 111(4):908—911.
, & R. Vargas. A new species of mud
shrimp, Upogebia cortesi, from Pacific Costa
Rica (Decapoda: Thalassinidea: Upogebi-
idae).—Proceedings of the Biological Society
of Washington 113(1):13—16.
2000b. Bishop, G. A., & . Fossil crabs from
Tepee Buttes, submarine seeps of the Late Cre-
taceous Pierre Shale, South Dakota and Colo-
rado, U.S.A.—Journal of Crustacean Biology
(Special no. 2):(in press).
In press. . Comment on the proposed designa-
tion of Scottia pseudobrowniana Kempf, 1971
as the type species of Scottia Brady & Norman,
1889 (Crustacea: Ostracoda) (Case 2896; see
BZN 51(4):304—305).—Bulletin of Zoological
Nomenclature.
In press. . Comment on the proposed conser-
vation of both the generic and specific names
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BZN 52(2):153—156).—Bulletin of Zoological
Nomenclature.
1997d.
1998b.
2000a.
List of taxa named by Austin B. Williams
Taxa are listed alphabetically within each
major decapod group. Holotype deposition
and number is indicated for all species and
subspecies. Asterisk indicates fossil taxa.
Abbreviations for repositories are as fol-
lows: AHKE Allan Hancock Foundation,
University of Southern California (now
Natural History Museum of Los Angeles
County); AMNH, American Museum of
Natural History, New York; MCZ, Museum
of Comparative Zoology, Harvard Univer-
sity, Cambridge, Massachusetts; MZUSP,
Museu de Zoologia, Universidade de Sao
Paulo, Brazil; SDSM, Museum of Geology,
South Dakota School of Mines, Rapid City;
SDSNH, San Diego Society of Natural His-
tory, California; UKMNH, University of
Kansas Museum of Natural History,
Lawrence; USNM, National Museum of
Natural History, Smithsonian Institution,
Washington, D.C.
Decapoda
Caridea
Alvinocaris Williams & Chace, 1982b.
Alvinocaris lusca Williams & Chace,
1982b. USNM 184534.
Alvinocaris markensis Williams, 1988c.
USNM 234286.
Alvinocaris muricola Williams, 1988c.
USNM 234288.
Alvinocaris stactophila Williams, 1988c.
USNM 234291.
Leptalpheus Williams, 1965b.
Leptalpheus forceps Williams, 1965b.
USNM 111084.
Ogyrides hayi Williams, 198la. USNM
47958.
Ogyrides limicola Williams, 1955a. USNM
96675.
Opaepele Williams & Dobbs, 1995c.
Opaepele loihi Williams & Dobbs, 1995c.
USNM 251447.
Rimicaris Williams & Rona, 1986c.
Rimicaris chacei Williams & Rona, 1986c.
USNM 228452.
Rimicaris exoculata Williams & Rona,
1986c. USNM 228443.
Astacidea
Homarinus Kornfield, Williams & Steneck,
1995b.
Orconectes eupunctus Williams, 1952a.
UKMNH T4250.
Orconectes meeki brevis Williams, 1952a.
UKMNH T8140.
Orconectes nana marcus Williams, 1952a.
UKMNH T4970.
Orconectes nana
UKMNH T6640.
Williams, 1952a.
VOLUME 113, NUMBER 1
Orconectes neglectus chaenodactylus Wil-
liams, 1952a. UKMNH T4420.
Orconectes ozarkae Williams,
UKMNH T6150.
LO524.
Thalassinidea
Aethogebia Williams, 1993b.
Aethogebia gorei Williams, 1993b. USNM
251425.
*Axiopsis eximia Kensley & Williams,
1990c. USNM 219431.
Calocaris (Calastacus) jenneri Williams,
1974b. USNM 150472.
Calocaris (Calastacus) oxypleura Williams,
1974b. USNM 101651.
Pomatogebia Williams & Ngoc-Ho, 1990e.
Upogebia acanthops Williams, 1986a.
USNM 213194.
Upogebia aestuari Williams, 1993b.
USNM 251407.
Upogebia aquilina Williams, 1993b.
USNM 251426.
Upogebia baldwini Williams, 1997d.
USNM 251486.
Upogebia bermudensis Williams, 1993b.
MCZ 12873.
Upogebia burkenroadi Williams, 1986a.
SDSNH 3985.
Upogebia careospina Williams, 1993b.
USNM 138899.
Upogebia casis Williams, 1993b. USNM
251224.
Upogebia cocosia Williams, 1986a. USNM
213268.
Upogebia coralliflora Williams & Scott,
1989b. USNM 230075.
Upogebia cortesi 2000a. USNM 291186.
Upogebia dawsoni Williams, 1986a.
AHF2566.
Upogebia felderi Williams, 1993b. USNM
251430.
Upogebia galapagensis Williams, 1986a.
USNM 213223.
Upogebia inomissa Williams,
USNM 251396.
Upogebia jonesi Williams, 1986a. USNM
213195.
1993b.
|
Upogebia lepta Williams, 1986a. USNM
213270.
Upogebia maccraryae Williams, 1986a.
USNM 213202.
Upogebia macginitieorum Williams, 1986a.
USNM 213219.
Upogebia molipollex Williams, 1993b.
AMNH 6820.
Upogebia omissago
USNM 222057.
Upogebia onychion Williams, 1986a. AHF
4133.
Upogebia paraffinis Williams,
MZUSP 8049
Upogebia_ pillsbury
USNM 251435.
Upogebia ramphula Williams,
USNM 213446.
Upogebia schmitti Williams, 1986a. AHF
3933:
Upogebia spinistipula Williams & Heard,
199la. USNM 239251.
Upogebia synagelas Williams,
USNM 233572.
Upogebia tenuipollex Williams, 1986a.
USNM 213236.
Upogebia thistlei Williams, 1986a. USNM
21S251%
Upogebia toralae Williams & Hernandez-
Aguilera, 1998b. USNM 285522.
Williams, 1993b.
1993b.
1993b.
Williams,
1986a.
1987d.
Upogebia vargasae Williams, 1997d.
USNM 251484.
Upogebia veleronis Williams, 1986a.
(USNM 213272)
Anomura
Munidopsis alvisca Williams, 1988c.
USNM 234294.
Munidopsis glabra Pequegnat & Williams,
1995e. USNM 251455.
Munidopsis granosicorium Williams &
Baba, 1990b. USNM 240205.
Munidopsis lentigo Williams & Van Dover,
1983b. USNM 191160.
Munidopsis lignaria Williams & Baba,
1990b. USNM 240202.
Munidopsis marianica Williams & Baba,
1990b. USNM 240198.
{[F9,
Shinkaia Baba & Williams, 1998a.
Shinkaia crosnieri Baba & Williams,
1998a. USNM 251480.
Uroptychus edisonicus Baba & Williams,
1998a. USNM 251479.
Brachyura
Allactaea Williams, 1974a.
Allactaea lithorostrata Williams,
USNM 143770.
Bothromaia Williams & Moffit, 1991b.
Bothromaia griffini Williams & Moffit,
1991b. USNM 250884.
Bythograeoidea Williams, 1980. (Super-
family).
Bythograeidae Williams, 1980. (Family).
Bythograea Williams, 1980.
Bythograea mesatlantica Williams, 1988c.
USNM 234300.
Bythograea thermydron Williams,
USNM 172830.
Callinectes similis Williams, 1966b. USNM
113341.
Cyclozodion Williams & Child, 1989a.
Cyclozodion tuberatum Williams & Child,
1989a. USNM 234462.
Epilobocera wetherbeei Rodriguez & Wil-
liams, 1995a. USNM 268832.
Eplumula Williams, 1982a.
*Heus Bishop & Williams, 2000b.
*Heus foersteri Bishop & Williams, 2000b.
SDSM 11016.
Hypsophrys noar Williams, 1974c. USNM
150816.
1974a.
1980.
Latreillia manningi Williams, 1982a.
USNM 57071.
Latreillia metanesa Williams, 1982a.
USNM 74570.
Menippe adina Williams & Felder, 1986e.
USNM 228862.
Mimilambridae Williams, 1979c. (Family).
Mimilambrus Williams, 1979c.
Mimilambrus wileyi Williams,
USNM 172222.
*Necrocarcinus olsonorum Bishop & Wil-
liams, 1991c. SDSM 11000.
19796.
2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Ovalipes stephensoni Williams, 1976a.
USNM 155110.
Panopeus austrobesus Williams, 1984a.
USNM 59462.
Panopeus margentus Williams & Boschi,
1990d. USNM 239191.
Panopeus meridionalis Williams,
USNM 99846.
*Plagiophthalmus bjorki Bishop & Wil-
liams, 2000b. SDSM 11021.
*Raninella manningi Bishop & Williams,
2000b. SDSM 11018.
Rochinia decipiata Williams & Eldredge,
1994. USNM 251434.
Stilbomastax Williams, Shaw & Hopkins,
1977a.
1984a.
Acknowledgments
We would like to thank David McN. Wil-
liams for providing valuable family facts,
and also several of Austin’s colleagues and
friends for contributing photographs and
other information. In particular, we would
like to mention Norma Samuels who kindly
provided photographs and insight on as-
pects of Austin private life; Gale A. Bishop,
for providing valuable and often emotional
testimony on his experiences in the field;
Raymond B. Manning and Brian Kensley
for reminiscing on their interactions with
Austin; and Keiji Baba for information and
photographs. The invaluable help of Mi-
chael Vecchione and the dedicated staff of
the Systematics Laboratory, National Ma-
rine Fisheries Service, at the Smithsonian,
in locating and compiling files and data on
Austin’s professional life, is gratefully ac-
knowledged; these include Lara Cooper,
Ruth Gibbons, David Hardy, Keiko Hira-
tsuka Moore, Roosevelt McMillan, Mary
Mickevich, Tom Munroe, and Martha Ni-
zinski. Thanks also to Rose A. Gulledge for
processing electronic images of the photo-
graphs, and compiling bibliographic infor-
mation; Chad Walter for proof-reading the
bibliography; and Molly K. Ryan for put-
ting together the figure.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):13—16. 2000.
A new species of mud shrimp, Upogebia cortesi, from Pacific Costa
Rica (Decapoda: Thalassinidea: Upogebiidae)
Austin B. Williams} and Rita Vargas
(ABW) National Marine Fisheries Service Systematics Laboratory, National Museum of
Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; (RV) Museo de
Zoologia, Escuela de Biologia, Universidad de Costa Rica, 2060 Costa Rica (+: deceased
October 27, 1999)
Abstract.—Upogebia cortesi, a new species of mud shrimp from the Pacific
side of Costa Rica is described and illustrated. The type series was dredged
parallel to shore in water of 30—40 m depth. The species shares lack of a strong
proximal mesioventral spine on the merus of the second pereopod with several
members of the genus from the western hemisphere; moreover, the species
stands alone in having spineless articles on pereopods 1—5 except for fingers
of the chelae. The rostrum has no ventral spines. Similarities between this and
related species are emphasized in a partial abridgment of the key to upogebiid
species in the eastern Pacific.
Eight species of the family Upogebiidae
have been reported for the Pacific coast of
Costa Rica (Vargas & Cortés 2000). Of
these Pomatogebia cocosia (Williams
1986) and Upogebia vargasae Williams,
1997, were described based on material
from Costa Rica. During the 1998 Mollusk
Workshop organized by INBio at the “‘Re-
serva Absoluta de Cabo Blanco’’, Peninsula
de Nicoya, a dredge sample conducted be-
tween the mainland and Isla Cabo Blanco,
at 30—40 m depth, yielded three specimens
of a new species of Upogebia Leach, 1814,
described here. The substrate where the
specimens were obtained included calcare-
ous algae and rock fragments; at least 10
species of brachyuran crabs and several
mollusk species were also found in the sam-
ple.
Specimens are deposited in the National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. (USNM), and
in Museo de Zoologia, Escuela de Biologia,
Universidad de Costa Rica, San José
(UCRMZ), as indicated in the species ac-
count.
Upogebia cortesi, new species
Fig. 1
Material.—Costa Rica, NW side of Isla
Cabo Blanco, dredged parallel to coast, 30—
40 m, 16-17 May 1998: USNM 291186,
male holotype; UCRMZ 2220-06, male par-
atype; USNM 291187, female ovigerous
paratype.
Diagnosis.—Rostrum with sides convex
in dorsal view, obsolescent pair of spines
on submedian anterior margin; projections
to either side of rostrum slender and spine-
like; no spine on postocular margin; ante-
rior gastric region bearing many spines
nearly hidden in patch of dense setae. Ab-
dominal sternites unarmed. Telson subrect-
angular, sides slightly crenulate. Merus of
cheliped lacking subdistal dorsal spine and
spines on ventral margin; carpus essentially
spineless; palms spineless. Pereopods 2—5
spineless; pereopod 2 without proximal me-
sioventral spine on merus.
Description.—Rostrum (Fig. la,b) hori-
zontal in lateral view with tip slightly ex-
ceeding eyestalks; convex in dorsal aspect,
with pair of submedian obsolescent blunted
14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig? tl.
Upogebia cortesi, new species, male holotype, USNM 291187: a, Carapace and cephalic region,
lateral; b, Anterior carapace, dorsal; c, Cheliped, right lateral; d, Chela and carpus, right mesial; e, Right pereopod
2; f, Right pereopod 3; g, Right pereopod 4; h, Right pereopod 5; i, Telson, left uropod, and part of abdominal
segment 6, dorsal. Scale = 3 mm.
spines on anterior margin; 2 subdistal erect
acute dorsal spines followed on each side
by about 6 smaller marginal spines along
either side of field of small spines on an-
terior gastric region of carapace, spines on
anterior part of field hidden in dense patch
of setae. Lateral ridge on either side of an-
terior gastric region extended anteriorly to
spine similar in size to dorsal spines on ros-
trum, ridge bearing crest of 18 spines, an-
terior 4 spines larger than succeeding
spines. Posterior region of carapace gla-
brous. Shoulder lateral to cervical groove
bearing no spines below intersection with
thalassinidean line, latter continuing to pos-
terior margin of carapace without interrup-
tion; postocular margin of carapace un-
armed.
Abdominal sternites unarmed.
Telson (Fig. 17) subrectangular, wider
than long, posterior margin nearly straight,
smooth; transverse proximal ridge promi-
VOLUME 113, NUMBER 1
nent, rather broad, lateral ridge at each side
also rather broad; lateral margins almost
imperceptibly crenulate.
Eyestalk stout, horizontal, shorter than
rostrum; lower margin slightly convex; cor-
nea narrower than diameter of stalk and di-
rected laterally.
Antennular peduncle (Fig. la) reaching
to about base of terminal article of antennal
peduncle, combined length of proximal 2
articles subequal to length of terminal arti-
cle.
Antennal peduncle (Fig. la) with distal
article and distal half of penultimate article
extending beyond tip of rostrum; moderate
compressed scale bearing obsolescent an-
teroventral spine.
Maxilliped 3 bearing epipod.
Epistomial projection rather broad in lat-
eral view, bearing prominent, acuminate
apical spine.
Chelipeds (Fig. lc,d) lacking spine on
ventral margin of merus. Carpus trigonal,
with barely perceptible longitudinal lateral
groove, obsolescent spine at anterior ven-
trolateral corner. Chela length about 2.7
times chela height. Fixed finger slender,
with extended tip. Dactyl thick, longer than
fixed finger, drawn to corneous tip, and at
midlength bearing small tooth on occlusive
edge opposing tip of fixed finger.
Pereopod 2 (Fig. le) reaching about to
distal edge of cheliped palm; pereopods 2-—
4 (Fig. 1f-/) spineless. Pereopod 5 of usual
form, with cleaning brush on propodus.
Uropods (Fig. 17) with acute spine on
protopod above base of mesial ramus; both
rami slightly exceeding telson, and with
distal margins smooth.
Measurements (in mm).—Male holotype,
anterior carapace length 6.1, carapace
length 9.1, length of chela including fixed
finger 4.1, mid-length height of chela 1.8;
male paratype, same, 4.8, 7.8, 3.8, 1.8; fe-
male ovigerous paratype, same, 5.4, 7.9,
42, 1:5:
Known range.—Known only from the
type locality.
Etymology.—The species is named for
15
Dr. Jorge Cortés, Curator of Cnidaria, Mu-
seo de Zoologia, Escuela de Biologia,
Universidad de Costa Rica, in recognition
of his life-long efforts to advance knowl-
edge of the marine fauna from Costa Rica.
Remarks.—Upogebia cortesi, new spe-
cies, shares with several eastern Pacific and
western Atlantic members of the genus a
second pereopod on which the merus bears
no proximal mesioventral spine (see keys to
species in Williams 1986, 1993). The ab-
dominal sternites and pleura bear no ventral
spinules.
The species stands alone, however, with
respect to several other characters. The ros-
trum with rounded anterior margin has no
ventral spines. The anterior gastric region
and rostrum bear an extremely dense patch
of setae obscuring spines on this surface.
The pereopods are virtually spineless. This
condition is most easily demonstrated by
emendation of the key to species of the
eastern Pacific (Williams 1986) in which
the comparative relationship to other spe-
cies in the region becomes apparent. Al-
though the new species has spineless pereo-
pods, it appears to be related to U. tenui-
pollex Williams, 1986 which has a well de-
veloped distodorsal spine on the merus of
the cheliped and a cluster of spines on the
merus of pereopod 3.
Emended part of Williams (1986:7—10)
key distinguishing U. cortesi, new species,
and U. tenuipollex
4. Merus of pereopod 2 lacking proximal
MESIOVERINAL SPINE. 6). cies own dee eens os 5
Merus of pereopod 2 bearing proximal
TUE STOMEMELON SOHNE gg 5p Sc a ch ig sr wenn 12
DS: Posltoctiar Spine @OSEML «6... 4s... > 6a
Postocular spine present and well de-
WER Cner teh reer ee Ss at oe 8
6a. Merus of cheliped and pereopod 3
SpINCIESS 7. PF, U. cortesi, new species
Merus of cheliped and pereopod with
SS). Were esa Re ee TI eal 6
6. Merus of chelipeds bearing well devel-
oped distodorsal spine; merus of pereo-
pod 3 bearing cluster of proximoventral
16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
SHINES 24.2)! Se. eee U. tenuipollex
Williams Merus of chelipeds lacking
distodorsal spine or with spine tiny;
merus of pereopod 3 bearing few ob-
solescent proximoventral spines .... 7
Acknowledgments
This study was made possible, in part, by
Short-Term Visitor grant awarded to RV by
the Office of Fellowships and Grants,
Smithsonian Institution, Washington, D.C.
We thank Michael Vecchione for critical
reading of the manuscript and Keiko Hira-
tsuka Moore for preparing the illustrations.
This is a contribution of the Museo de
Zoologia, Escuela de Biologia, Universidad
de Costa Rica.
Literature Cited
Leach, W. E. 1814. Crustaceology. /n Edinburgh En-
cyclopaedia, 2nd edition, 7:383—437, plate 221.
New York: Samuel Whiting and John L. Tiffany.
Vargas, R., & J. Cortés. 2000. Biodiversidad marina
de Costa Rica: Crustacea: Decapoda (Penaeo-
idea, Sergestoidea, Caridea, Astacidea, Thalas-
sinidea, Palinura) del Pacifico.—Revista de
Biologia Tropical 47 (in press).
Williams, A. B. 1986. Upogebia, from the eastern Pa-
cific (Thalassinoidea: Upogebiidae)—Memoir
of the San Diego Society of Natural History 14:
1-60.
. 1993. Mud shrimps, Upogebiidae, from the
western Atlantic (Crustacea: Decapoda: Thal-
assinidae).—Smithsonian Contributions to Zo-
ology 544:1-77.
. 1997. Two new species and a range extension
of mud shrimps, Upogebia from Pacific Costa
Rica and Mexico (Decapoda: Thalassinidea:
Upogebiidae).—Proceedings of the Biological
Society of Washington 110:617—623.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):17—23. 2000.
Periclimenes murcielagensis, a new species of shrimp
(Crustacea: Decapoda: Palaemonidae) living on black coral from the
Pacific coast of Costa Rica
Rita Vargas
Escuela de Biologia, Universidad de Costa Rica, 2060 San José, Costa Rica
Abstract.—Periclimenes murcielagensis, new species, a pontoniine shrimp
living on black coral colonies, occurs at depths of 25 m at Isla San Pedrito,
Archipiélago de las Islas Murciélago, Guanacaste, Costa Rica. This new species
is most similar to P. veleronis, Holthuis, and P. americanus Kingsley, and is
distinguished from these by characters from the rostrum, basal antennular seg-
ment, antennular flagellum, and incisor process of the mandible.
Shrimps of the genus Periclimenes Cos-
ta, 1844, belong to the subfamily Pontoni-
ine, and are distributed widely in subtropi-
cal and tropical marine waters worldwide.
Some species live freely but the majority
are associated with other marine inverte-
brates. Of the species of this genus known
from the eastern Pacific, only P. infraspinis
(Rathbun 1902), P. lucasi Chace, 1937, and
P. veleronis Holthuis, 1951, have not been
reported in association with other inverte-
brates (Holthuis 1951). Known associates
of these shrimps in the western Atlantic are
sponges, gorgonians, actinians, corallimor-
pharians, rhizostome scyphozoans, hy-
droids, antipatharians, bivalves, ophiuroids
and crinoids (Heard & Spotte 1991, Spotte
et al. 1994).
While conducting ecological studies of
soft corals, specimens of an undescribed
species of Periclimenes were found living
on colonies of black coral (Antipathes pan-
amensis Verrill, 1869) at Isla San Pedrito,
Archipiélago de las Islas Murciélago,
Guanacaste, Costa Rica. This new species
is described herein.
Specimens were collected during SCU-
BA dives at 25 m. Plastic bags were placed
over black coral colonies and closed with a
rubber band. The shrimps and Antipathes
panamensis colonies were fixed in a mix-
ture of 10% formalin and seawater, the col-
onies washed in the laboratory with fresh
water, and all the liquid passed through a
0.5 mm mesh. The shrimps were preserved
in 70% ethanol.
The material is deposited in the National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. (USNM), and
Museo de Zoologia, Universidad de Costa
Rica (UCRMZ). Carapace length (CL) was
measured from the tip of the rostrum to the
posterodorsal margin of the carapace.
Periclimenes murcielagensis, new species
Figs. 1-5
Material.—Holotype: ovigerous female
(CL 7.75 mm), USNM 260931, Isla San
Pedrito, Archipiélago de las Islas Murcié-
lago, Guanacaste, Costa Rica, 25 m, coll.
Odalisca Breedy, 24 May 1996.—Para-
types: 4 females (CL 7.5—7.9 mm), 2 males
(CL 5.256.5), USNM 260932; 9 females
(CL 6.25—-8.4 mm), 2 males (CL 5.6—6.25
mm), UCRMZ 2247-01, same collection
data as holotype.
Diagnosis.—Carapace armed with both
hepatic and antennal spine; rostrum well
developed, reaching end of antennular pe-
duncle, usually armed with 8—10 teeth dor-
sally and 2—4 teeth on distal half of ventral
margin. Antennular peduncle having basal
18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
yes BESS
— oe
= ————
—— = ——
eS oS Se ee as Ss ee SS
=
an
e,
2
v
SB
Ss
eee
a
a = oe
Periclimenes murcielagensis, new species, holotype ovigerous female (CL 7.75 mm), lateral view
USNM 260931. Left antennule, antenna and pereopods 1—5 not shown. Scale equals 5 mm.
article armed with 2—4 distolateral spines.
Pereopods 3—5 with dactylus simple.
Description.—Rostrum (Figs. 1, 5a—f)
nearly straight, slightly reaching end of an-
tennular peduncle; dorsal margin armed
with 8—10 teeth interspaced with setae, pos-
teriormost tooth placed on carapace poste-
rior to level of orbital margin, second tooth
on carapace usually even with, or anterior
to, orbital margin. Distance between first
and second tooth, sometimes slightly larger
than distance between other teeth, which
are regularly spaced over rostrum; ventral
margin slightly convex, armed with 2-4
teeth on distal half. No postorbital ridges or
supra orbital spines. Antennal spine well
developed, acute. Hepatic spine well devel-
oped, slightly larger and more robust than
antennal spine. Lower orbital angle pro-
duced into small, blunt lobe.
Abdominal pleura broadly rounded.
Sixth somite (Fig. 1) nearly twice as long
as fifth and slightly longer than telson. Dor-
sal spines of telson (Fig. 2c) distinct, prox-
imal pair situated near midlength of telson,
distal pair of spines closer to proximal pair
than posterior margin of telson; intermedi-
ate marginal spines at posterior end of tel-
son not quite twice as long as mesial pair.
Cornea as broad as peduncle (eyestalk),
constricted at junction with eyestalk, acces-
sory pigment spot and associated ommatid-
ia present on dorsoproximal margin of cor-
nea.
Antennular peduncle (Figs. 2b, 4g—h)
with stylocerite sharp and slender, reaching
to about midlength of basal segment; dis-
tolateral margin of basal segment armed
with 2—4 spines, second and third segment
subequal in length and width. Lateral an-
tennular flagellum with 2 branches fused
for about 8 joints; portion of shorter branch
not fused consisting of about 6 joints, and
about 0.5 times shorter than fused portion.
Antennal scale (Fig. 2a) reaching distal
margin of third antennular segment, more
than 3 times as long as broad; lateral mar-
gin nearly straight, distal tooth falling far
short of strongly produced anteromesial an-
gle of blade. Antennal peduncle reaching
about to midlength of scale; basal segment
with sharp lateral spine near base of scale.
Mouthparts as figured (Fig. 3a—f). Man-
dible (Fig. 3a) lacking palp; incisor process
ending in 4 distinct teeth, distal tooth larg-
est; molar process dentate. Maxilla 1 (Fig.
3b) with upper endite (lacinia) possesing
stout apical spine-setae crown, distal to, 10
or more smaller subapical setae; endite with
4 or more stout apical spine-setae and 6 or
more subapical setae on each side. Maxilla
2 (Fig. 3c) with entire endite. Maxilliped 1
VOLUME 113, NUMBER 1
4
Fig. 2. Periclimenes murcielagensis, new species. Paratype male (CL 5.6 mm) UCRMZ 2247-01. a, anterior
part of carapace; b, right antennule; c, telson; d, appendix masculina. Scale equal 2 mm (a, c), 1 mm (b), 0.25
mm (qd).
(Fig. 3d) with well developed exopodal fla-
gellum (lash) possessing 4 terminal plu-
mose spine-setae; epipod slightly bilobed;
palp slender and laching terminal spine se-
tae. Maxilliped 2 (Fig. 3e) possessing well
developed exopod with terminal plumose
setae; epipod rectangular. Maxilliped 3
(Fig. 3f) extending for proximal 0.25 of
scaphocerite; exopod exceeding midlength
of proximal segment; with broad, round
epipod.
First pereopod (Fig. 4a) reaching end of
antennal scale; fingers unarmed, shorter
than palm; carpus distinctly longer than
chela, subequal to merus. Second pereopod
distinctly unequal. Major cheliped (Fig. 4b,
Cc) overreaching antennal scale by approxi-
mately length of chela; fingers armed with
teeth, distinctly shorter than palm, carpus
distinctly shorter than chela, subequal to
merus; ischium and merus equal in length.
Minor cheliped (Fig. 4d) of second pair
overreaching antennular scale by approxi-
mately length of fingers; fingers unarmed,
shorter than palm; chela, carpus, merus and
ischium subequal in length. Pereopods 3—5
(Fig. 4e—g) nearly equal in size and shape;
dactyls entire (not bifid), propodi with sin-
gle spine on distal flexor margin. Third pe-
reopod just reaching end of antennal scale;
propodus 4 times length of dactyl, slightly
more than twice length of carpus or ischi-
um; merus subequal in length to propodus,
with single distal spine on flexor margin.
Fourth pereopod extending to distal end of
second segment of antennular peduncle;
propodus 4 times length of dactyl, slightly
more than twice length of carpus or ischi-
um; merus subequal in length to propodus.
Fifth pereopod extending to distal end of
second segment of antennular peduncle;
propodus 4 times length of dactyl, subequal
20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Big. 3:
dible; b, maxilla 1; c, maxilla 2; d, maxilliped 1; e, maxilliped 2; f, maxilliped 3. Scale equal 0.5 mm (a-e), 1
mm (f).
to merus; carpus and ischium subequal in
length.
Males with appendix masculina (Fig. 2d)
armed with 3 apical spine-like setae, midle
smaller and weakly serrate, and subapical
spine-like setae. Eggs size ranging in max-
imum length from 0.3 to 0.5 mm (hatching
Stage).
Uropodal exopods extending beyond tel-
son for about 0.8 times length of exopod;
with strong movable spine between disto-
lateral tooth and blade; movable spine dis-
tinctly longer than distolateral spine.
Color.—Orange after preservation.
Habitat.—Periclimenes murcielagensis,
new species, was found living on colonies
Fe
> N
cs y~
:
Periclimenes murcielagensis, new species. Paratype male (CL 5.6 mm) UCRMZ 2247-01. a, man-
of black coral Antipathes panamensis. Also
found on the same colonies with the pon-
toniin Waldola schmitti Holthuis, 1951, cir-
ripeds, mollusks and polychaete worms.
Depth: 25 m.
Distribution.—Known only from type lo-
cality, Archipiélago de las Islas Murciélago,
Guanacaste, Costa Rica.
Etymology.—The species is named for
the type locality, Archipiélago de las Islas
Murciélago.
Remarks.—Adult males and females
differ only in the size of the major cheli-
ped of the second pereopod; in males the
major cheliped is markedly smaller than
in females. The major cheliped in adult
VOLUME 113, NUMBER 1 21
G
é€
rr
Wf
Fig. 4. Periclimenes murcielagensis, new species. Paratype male (CL 5.6 mm), lateral view, UCRMZ 2247-
01. a, distal portion of chela of first pereopod; b, right second major pereopod; c, enlargement of chela of mayor
pereopod; d, distal portion of chela of minor pereopod; e, f, g, third, fourth and fifth pereopods. Scale equal 4
mm (b), 2 mm. (e, f, g), and 1 mm (a, c, d).
males is similar to that of immature fe- cies, is most similar to Periclimenes vele-
males. ronis, from La Libertad, Ecuador (Holthuis
Among the eastern Pacific species of 1951). The new species can be distin-
Periclimenes, P. murcielagensis, new spe- guished from P. veleronis by the slender
22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
7 oe 4
+ ae
ge oe
yf
—_———S Ss
Fig. 5.
Periclimenes murcielagensis, new species. Variations 1n rostrum (a—f) and basal segment of antennular
peduncle (g—j). (a, f, males; b, c, females; g, i, j, females; h, male). Scale equals 5 mm.
shape and number of teeth on the dorsal (8—
10) and ventral (2—4) margins of the ros-
trum; the presence of two to four subequal
spines on the distolateral margin of basal
segment of antennular peduncle; the two
branches of the antennular flagellum are
fused for about eight joints (five joints in
P. veleronis); the non-fused portion consists
of about six joints and is about one-half
times shorter than fused portion, whereas in
P. veleronis the non-fused portion has three
joints and is more than half as long as the
fused portion.
Periclimenes murcielagensis, new spe-
cies, also resembles P. americanus, from
Florida (Kinsgley 1878). The two have a
similarly shaped and armed rostrum. The
new species can be distinguished from P.
americanus by the presence of two to four
subequal spines on the distolateral margin
of the basal segment of antennular peduncle
(one in P. americanus); no postorbital ridge
is present in P. murcielagensis; the two
branches of the antennular flagellum are
fused for about eight joints (eight to 12
joints in P. americanus), and the non-fused
portion consists of about six joints (three or
four in P. americanus); the incisor process
of the mandible ends in four distinct teeth
in P. murcielagensis (three in P. american-
us); the second pereopods are distinctly un-
equal in P. murcielagensis (equal in P.
americanus).
Acknowledgments
This study was possible thanks to a
Short-Term Visitor award, granted by the
VOLUME 113, NUMBER 1
Office of Fellowships and Grants, Smith-
sonian Institution, Washington, D.C. I thank
““Area de ConservaciOn Guanacaste, Min-
isterio del Ambiente y Energia’”’ for arrang-
ing a visit to Islas Murciélago. I also thank
R. Lemaitre and R. Heard for their critical
comments on the manuscript; J. Cortés for
his constant support, and O. Breedy for
sending me the specimens. This is a contri-
bution of the Museo de Zoologia, Escuela
de Biologia, Universidad de Costa Rica.
Literature cited
Chace, FE A., Jr. 1937. Caridean decapod crustacea
from the Gulf of California and the west coast
of Lower California. Part VII. The Templeton
Crocker Expedition.—Zoologica (New York)
22:109-138.
Costa, O. G. 1844. Su due nuovi generi di Crostacei
Decapodi Macrouri Nota.—Annali delle Acca-
demia degli Aspiranti Naturalisti, Napoli 2:
290-291.
Heard, R. W., & S. Spotte. 1991. Pontoniinae shrimps
(Decapoda: Caridea: Palaemonidae) of the
northwest Atlantic. II. Periclimenes patae new
23
species, a gorgonian associate from shallow reef
areas off Turks and Caicos Islands and Florida
Keys.—Bulletin of Marine Sciences 55(1):212
DLT.
Holthuis, L. B. 1951. A general revision of the Palae-
monidae (Crustacea Decapoda Natantia) of the
Americas. I. The subfamilies Euryrhynchinae
and Pontoniinae.—Occasional Papers, Allan
Hancock Foundation Publications 11:1—331.
Kingsley, J. S. 1878. Notes on the North American
Caridea in the Museum of the Peabody Acad-
emy of Science at Salem, Mass.—Proceedings
of the Academy of Natural Sciences of Phila-
delphia 1878:89-98.
Rathbun, M. J. 1902. Description of new decapod crus-
taceans from the west coast of North Ameri-
ca.—Proceeding of United States National Mu-
seum 24:885-—905.
Spotte, S., R. Heard, & P. Bubucis. 1994. Pontoniinae
shrimps (Decapoda: Caridea: Palaemonidae) of
the northwest Atlantic. IV. Periclimenes anti-
pathophilus new species, a black coral associate
from the Turks and Caicos Islands and eastern
Honduras.—Bulletin of Marine Sciences 55(1):
212-227.
Verrill, A. E. 1869. Review of the corals and polyps
of the west coast of America.—Transactions of
Connecticut Academy of Science 1:377—558.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):24—29. 2000.
A new squat lobster of the genus Munidopsis Whiteaves, 1874
(Crustacea: Decapoda: Galatheidae) from Taiwan
Ming-Feng Wu and Tin-Yam Chan
Institute of Marine Biology, National Taiwan Ocean University, Keelung, Taiwan, R.O.C.
Abstract.—A new species of the squat lobster of the genus Munidopsis Whi-
teaves, M. formosa, is described from deep-water off the northeastern coast of
Taiwan. The new species is most similar to M. camelus Ortmann, but differs
in having a much broader and less curved rostrum, the posterior carapace ridge
generally armed with a pair of submedian spines, only the chelipeds bearing
epipods, and having a lighter coloration. The relationships of this new species
with some other similar species are also discussed.
There are about 150 known species of
Munidopsis Whiteaves, 1874 (see Baba
1988), most of which inhabit deep waters
(to 5330 m deep) and generally have small
eyes as well as a triangular rostrum. To
date, only three species, namely M. anda-
manica MacGilchrist, 1905, M. cylindro-
phthalma (Alcock, 1894) and M. latimana
Miyake & Baba, 1966, of this genus have
been reported from Taiwan (Wu et al.
1998). Recently, several specimens of an
undescribed species of Munidopsis from
Taiwan were collected off the northeastern
coast at a depth of about 500 m. Careful
examinations of these specimens showed
that they are most similar to M. camelus
Ortmann, 1892 from Japan, and differ from
other members of the genus in the rostrum
being trifid, as well as the second and the
third abdominal tergites each armed only
with a pair of large submedian spines. De-
tailed comparison with M. camelus revealed
several slight but constant differences be-
tween the Taiwanese and Japanese material.
The Taiwanese specimens represent a new
species described herein.
Specimens of the new species are depos-
ited in the National Taiwan Ocean Univer-
sity, Keelung (NTOU), and those used of
M. camelus in the personal collection of
Prof. K. Baba, Kumamoto University, Ja-
pan (KB). The measurements given are of
carapace length (cl) excluding rostrum.
Munidopsis formosa, new species.
Figs. -la,.c=e, (2a. ¢. .d.5
Material examined.—Holotype: Taiwan,
northeastern coast, Tai-Shi fishing port, I-
Lan County, commercial trawlers, about
500 m, soft bottoms, Aug 1998, 1 6, cl
20.4 mm (NTOU-H 1998-08).
Paratypes: Taiwan, northeastern coast,
Tai-Shi fishing port, I-Lan County, com-
mercial trawlers, about 500 m, soft bottoms,
Apr 1997, 1 6, cl 27.8 mm (NTOU-P
1997-04); 15 May 1998, 1 6, cl 25.3 mm,
1 ovigerous 2, cl 22.5 mm (NTOU-P 1998-
05-15); 28 Apr 1999, 1 ovigerous @, cl
17.4 mm (NTOU-P 1999-04-28).
Description.—Body entirely covered
with fine short setae. Rostrum broad, about
2.5 times as long as wide and % as long as
carapace; more or less horizontal, with tip
gently curving dorsad; carinate dorsally,
with trifid tip. Carapace (Figs. la, 2a)
slightly longer than wide; frontal margin
with 1 spine between rostrum and antero-
lateral spines; lateral margin feebly convex,
armed with 4 stout spines (including an-
terolateral spine) at distal half; 1 pair of epi-
gastric spines present; gastric region mod-
erately convex, bearing row of 1-3 longi-
VOLUME 113, NUMBER 1
tudinal spines; cervical region moderately
excavated; cardiac region raised, bearing
large median spine; posterior transverse
ridge elevated and generally armed only
with pair of large submedian spines.
Abdomen (Fig. 1a) with second and third
tergites each having pair of large subme-
dian spines. Telson (Fig. 2d) subdivided
into 10 plates; lateral margins fringed with
dense setae (very thick in male), posterior
margin with plumose setae.
Eyes small, lacking dark pigments, mov-
able, extending to about middle of rostrum.
A large spine present between eye and an-
tenna. Basal antennular segment (Fig. Ic)
bearing 2 strong distolateral spines, with
distal one slightly larger. Antennal peduncle
(Fig. 1d) with basal segment bearing dis-
tomesial and distolateral spines (distomesial
one very strong and long), other segments
spineless.
Merus of third maxilliped (Fig. le) lon-
ger than ischium, flexor margin bearing 3
spines, diminishing in size anteriorly, ex-
tensor margin armed with large distal spine.
Third thoracic sternite narrowing poste-
riorly, anterior margin concave; fourth tho-
racic sternite much wider, about 3 times as
wide as preceding (Fig. 2c).
Chelipeds subequal, long and robust
(massive in largest male), surface covered
with long setae; merus long and with some
large spines; carpus short, with spines
mainly restricted at distal margin; palm lon-
ger than finger and without large spines (but
sharply granular in the largest male); fin-
gers not perfectly gaping, inner margins
bearing some intermeshing teeth, outer
margins nearly straight except at tips (that
of fixed finger becoming convex in largest
male).
Walking legs robust, similar, all covered
with setae. First walking leg having merus
with large distodorsal and distolateral
spines, and some dorsal spines; carpus gen-
erally armed with 2 distodorsal spines; pro-
podus nearly straight, more than 5 times as
long as wide; dactylus much shorter than
25
propodus, distally curving ventrad, ventral
margin minutely dentate.
Epipod present only on chelipeds.
Eggs subspherical, about 0.5 mm in di-
ameter.
Coloration (Fig. 3).—Body pale orange
to orange, with color of females generally
deeper. Rostrum except tip, pale orange or
whitish. Fourth or fifth abdominal segments
to tailfan from pale orange to pale white
posteriorly. Eyes pale orange or nearly
whitish. Antennules same color as body but
antennal flagella orange red. Cervical
groove and cardiac depressions sometimes
whitish. Ventral surface whitish except
mouth parts and chelipeds pale orange.
Size.—Largest male and female cl 27.8
mm and 24.6 mm respectively. Smallest
ovigerous female cl 17.4 mm.
Distribution.—So far known only from
the northeastern coast of Taiwan, at depths
of about 500 m.
Type locality.—Taiwan, northeastern
coast.
Remarks.—The present form is closely
related to Munidopsis camelus Ortmann,
1892 from Japan and can be readily sepa-
rately from the other species of the genus
by the rostrum being trifid, as well as the
second and third abdominal tergites each
armed with a pair of large submedian
spines. Careful comparisons between M.
camelas (1.6 ¢ch23.2 mm. 1-2. cl 21.5 mm,
off Hayama, Sagami Bay, Japan, lobster
pot, Oct 1987, H. Ikeda coll. deposited at
Kumamoto University Faculty of Educa-
tion; also see Miyake & Baba 1967) and the
Taiwanese material revealed the following
differences. The anterior three pereiopods
bear distinct epipods in the Japanese ma-
terial but usually only the chelipeds have
epipods in the Taiwanese specimens. How-
ever, in one of the Taiwanese specimens
(NTOU-P 1997-04), a distinct epipod is
also present on the left first walking leg. It
seems that the presence or absence of epi-
pod are not always consistent as previously
thought for galatheids. Nevertheless, further
differences between the Taiwanese and Jap-
26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
PU UES
Wiad At sidatili
\s! anes IV, Ga ara
qu’
\
r { {
ta
= me ety j
acs OO Sa aii
= ii
AUT si 1)
Cty 1) galley
i iy
bay
Uy
4 iy i
Leh
Fig. 1. a, c-e: Munidopsis formosa, new species, d holotype cl 20.4 mm, N. E. Taiwan (NTOU-H 1998-
08). b: M. camelus Ortmann, 1892, 3d cl 23.2 mm, Sagami Bay, Japan (KB). a, carapace and anterior abdominal
somites, dorsal view; b, carapace; c, left basal antennular segment, ventral view; d, left antennal peduncle,
ventral view; e, basal segments of endopod of right maxilliped, ventral view. Scale bars = 5 mm.
VOLUME 113, NUMBER 1 27
Fig. 2. a, c—d: Munidopsis formosa, new species, 6 holotype cl 20.4 mm, N. E. Taiwan (NTOU-H 1998-
08). b: M. camelus Ortmann, 1892, d cl 23.2 mm, Sagami Bay, Japan (KB). a—b, carapace, lateral view; c,
anterior part of thoracic sternum, ventral view; d, telson, dorsal view. Scale bar = 5 mm.
anese material can be found. The posterior In the Taiwanese form, the posterior cara-
carapace ridge always bear a large median pace ridge generally armed with a pair of
spine which is accompanied with several submedian spines only (except in one spec-
large lateral spines in M. camelus (Fig. 1b). imen, NTOU-P 1999-1-19, there is one
28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Bige 3:
05-15).
more small spine present between the large
submedian spines) with the lateral parts al-
ways devoid of spines (Fig. la). Moreover,
the rostrum is distinctly narrower (near 4
times as long as wide) and with the tip
abruptly curving upward in the Japanese
material (Fig. 2b). The rostrum in the Tai-
wanese form, however, is rather broad (2.5
times as long as wide) and only gently
bending upward at tip (Fig. 2a). The col-
oration of the Japanese material also ap-
pears to be more reddish. Color photo-
graphs of a live specimen collected off
Boso Penisula (180—250 m) showed that
the body color of the Japanese material is
Munidopsis formosa, new species, ovigerous 2 paratype cl 22.5 mm, N. E. Taiwan (NTOU-P 1998-
orange-red. All the above differences show
that the Taiwanese form is distinct from M.
camelus and it is hereby described as new.
The present new species is also similar
to M. regia Alcock & Anderson, 1894, and
M. plumatisetigera Baba, 1988. However,
M. formosa new species, can be readily dis-
tinguished from M. plumatisetigera by the
epipods on the chelipeds and the less spiny
body. M. regia differs considerably from M.
formosa in having a much narrower and
longer rostrum, and a different spination on
the abdomen. Moreover, Alcock (1901)
mentioned that the color in life of M. regia
is “‘chalky pink’’.
VOLUME 113, NUMBER 1
Munidopsis formosa was collected from
about 500 m deep. This is probably the
main reason for this species being found
only recently. The fishing depth of local
deep-water trawlers have extended their
trawling depths down to 500—600 m, and
many deep-sea animals unknown to Taiwan
have been collected, including the present
new species.
Etymology.—This species is named after
its type-locality Taiwan since it is so far
only known from there. Formosa was the
old name of Taiwan and is used here as a
noun in apposition.
Acknowledgments
Grateful acknowledgment is extended to
Dr. K. Baba of the Kumamoto University
Faculty of Education for sending us on loan
the specimens of M. camelus and providing
us with many valuable comments; Dr. T.
Komai of Natural History Museum and In-
stitute, Chiba, for sending us color photo-
graphs of M. camelus, and S. H. Wu of our
laboratory for collecting most of the present
specimens. This work is supported by a re-
search grant from the National Science
Council, Taiwan, R.O.C.
Literature Cited
Alcock, A. 1894. Natural history notes from H. M.
Indian Marine Survey Steamer “Investigator,”
commander R. F Hoskyn, R. N., commaning.
Series II, no. 1. On the results of deep-sea
dredging during the season 1890-91 (contin-
ued).—Annals and Magazine of Natural Histo-
ry, series 6, 13:321—334.
. 1901. A descriptive catalogue of the India
deep-sea Crustacea Decapoda, Macrura and An-
29
omala in the Indian Museum. Being a revised
account of the deep-sea species collected by the
Royal Indian Marine Survey Ship Investigator.
Indian Museum, Calcutta. 286 pp.
, & A. R. S. Anderson. 1894. Natural history
notes from H. M. Indian Marines Survey
Steamer “Investigator”, commander C. F. Old-
ham, R. N., Commanding. Series I, No. 14. An
account of a recent collection of deep sea Crus-
tacea from the Bay of Bengal and Laccadive
Sea.—Journal of Asiatic Society of Bengal, 63
(II:3):141-185.
Baba, K. 1988. Chirostylid and Galatheid Crustacean
(Decapoda: Anomura) of the “‘Albatross”’ Phil-
ippine Expedition, 1907—1910.—Researches on
Crustacea, spec. no. 2:1—203.
MacGilchrist, A. C. 1905. Natural history notes from
the R. I. M.S. “Investigator’’. Capt. T. H. Hem-
ing, R. N. (retired), commanding. Series III. no.
6. An account of the new and some of the rarer
decapod Crustacea obtained during the survey-
ing seasons 1901—1904.—Annals and Magazine
of Natural History, series 7, 15:233-268.
Miyake, S., & K. Baba. 1966. Two new species of the
family Galatheidae from the Tosa Bay, Japan.—
Journal of the Faculty of Agriculture, Kyushu
University, Fukuoka 14(1):81—88.
Wis4 . 1967. New and rare species of the
family Galatheidae (Crustacea, Anomura) from
the Sagami Bay in the collection of the biolog-
ical Laboratory, Imperial Household, Japan.—
Journal of the Faculty of Agriculture, Kyushu
University 14(2):213-—224.
Ortmann, A. 1892. Die Decapoden-Krebse des Strass-
burger Museums. IV. Die Abtheilungen Galath-
eidea und Paguridea.—Zoologischen Jahrbuch-
ern, Abtheilung fiir Systematik, Geographie und
Biologie der Tiere, 6:241—326.
Whiteaves, J. EF 1874. On the recent deep-sea dredging
operations in the Gulf of St. Lawrence.—Amer-
ican Journal of Science (3)7:210—219.
Wu, M.F,.T. Y¥.Chan, & TH PB Yu. 1998.’On the Chi-
rostylidae and Galatheidae (Crustacea: Deca-
poda: Anomura) of Taiwan.—Annual of Taiwan
Museum 40:75—153. [In Chinese, with English
abstract]
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):30—38. 2000.
A new freshwater crab of the genus Geothelphusa Stimpson, 1858
(Crustacea: Decapoda: Brachyura: Potamidae) from Yakushima
Island, southern Kyushu, Japan
Hiroshi Suzuki and Tomokazu Okano
Marine Biological Laboratory, Faculty of Fisheries, Kagoshima University, 4-50-20 Shimoarata,
Kagoshima 890-0056, Japan
Abstract.—A new freshwater crab, Geothelphusa marmorata, is described
from Yakushima Island of Kagoshima Prefecture, southern Kyushu, Japan. It
is differentiated from congeners by possession of distally narrowed eyes, choc-
olate brown or dark red carapace with scattered black speckles in life, stout
gastric cristae, a deep H-shaped median gastro-cardiac depression, and laterally
curved penultimate segment of the male first gonopod with a mesially curved
ultimate segment. This is the second species of Geothelphusa Stimpson known
from Yakushima Island, and the twelfth species from Japan. Electrophoretic
analysis of 15 gene loci suggests that G. marmorata, new species, G. exigua
Suzuki & Tsuda and G. dehaani (White) are reproductively isolated.
Freshwater crabs of the genus Geothel-
phusa Stimpson, 1858 are distributed from
Taiwan, through the Ryukyu Islands, to the
Japanese mainland. Recently, a large num-
ber of species were reported from Taiwan
and Ryukyu Islands, and 39 species are
now recognized in the genus (Shy et al.
1994, Shy & Ng 1998, Tan & Liu 1998).
In Japan, 11 species are currently known,
1.e., G. dehaani (White, 1847), G. obtusipes
Stimpson, 1858, G. sakamotoana (Rathbun
1905), G. aramotoi Minei, 1973, G. tenui-
mana (Miyake & Minei 1965), G. levicer-
vix (Rathbun 1898), G. candidiensis Bott,
1970, G. miyazakii (Miyake & Chiu 1965),
G. exigua Suzuki & Tsuda, 1994, G. shok-
itai Shy & Ng, 1998, and G. minei Shy &
Ng, 1998 (see also de Haan 1835, Rathbun
1904, Bott 1967, Minei 1974b, Shy et al.
1994). Of these, G. dehaani is widely dis-
tributed on the Japanese mainland (north of
Honshu southward to Nakano-shima of the
Tokara Islands, south of Kyushu). The other
10 species are restricted to the southern
Kyushu or the Ryukyu Islands, including
Amami-ohshima.
During our current study of the popula-
tion genetics and geographic distribution of
G. dehaani and G. exigua in southern Kyu-
shu, unusual specimens of Geothelphusa
species were found on Yakushima Island in
Kagoshima Prefecture. The unusual eyes,
coloration of body, and structure of the
male first gonopods of these crabs indicate
that they represent a new species that is
herein described and illustrated. In addition,
a genetic analysis based on electrophoresis
is included.
Materials and Methods
For electrophoretic analysis, a total of
201 specimens of G. dehaani were collect-
ed from three populations (174 specimens
from Kotsuki River of the Kagoshima
mainland, 20 from Hitotsutani River of the
Kagoshima mainland, and seven from An-
boh River of the Yakushima Island); 40
specimens of G. exigua were collected from
two populations (20 specimens from Hitot-
sutani River and 20 from Kamiharai River
of the Kagoshima mainland); and 17 spec-
imens of the new species were collected
VOLUME 113, NUMBER 1
3]
Table 1.—List of enzymes and protein, and buffer systems used in electrophoretic analysis. CAPM 6.0; Citric
acid-aminopropyl morpholine, pH 6.0: CAPM 7.0; Citric acid-aminopropy! morpholine, pH 7.0: and CT 8.0;
Tris-citric acid, pH 8.0.
Enzyme and protein Symbol
(Abbreviation and E. C. number) for locus Buffer system
Aspartate aminotransferase (AAT, 2.6.1.1) AAT* CAPM 6.0
Adenylate kinase (AK, 2.7.4.3) AK* CAPM 7.0
Glyceraldehyde-3-phosphate dehydrogenase (GAPDH, 1.2.1.12) GAPDH* CAPM 6.0
Glucose-6-phosphate isomerase (GPI, 5.3.1.9) Grr CAPM 6.0, CAPM 7.0
Hexokinase (HK, 2.7.1.1) HkK= CTr-s:0
Isocitrate dehydrogenase (IDHP, 1.1.1.42) IDHP-I* CAPM 7.0
IDHP-2* CAPM 7.0
Lactate dehydrogenase (LDH, 1.1.1.27) IBD) « buy Cr sv
Malate dehydrogenase (MDH, 1.1.1.37) MDH- 1* CAPM 7.0
MDH-2* CAPM 7.0
Malic enzyme (ME, 1.1.1.38) ME* Cr s.6
Mannose-6-phosphate isomerase (MPI, 5.3.1.8) MPI* CT-E:0
Phosphogluconate dehydrogenase (PGDH, 1.1.1.44) PGDH* CAPM 7.0
Phosphoglucomutase (PGM, 5.4.2.2) PGM* CT 3.0
General protein (PROT) PROT* CAPM 7.0
from the Anboh River during 1997. Speci-
mens used for electrophoretic analysis were
stored at —35°C with a small volume of
freshwater. Muscles were extracted and ho-
mogenized with an equal volume of cold
distilled water. Horizontal starch gel elec-
trophoresis was performed for the detection
of enzyme and protein variations (Table 1).
Locus and gene nomenclature follows that
of Shaklee et al. (1990). Multiple loci for a
given enzyme were distinguished by nu-
merals, with “‘—/*’’ representing the most
anodally-migrating isozyme. All alleles
studied are designated alphabetically. The
genetic distance was calculated using Nei’s
formula (Nei 1972). All the specimens were
collected by the junior author.
The holotype and a paratype are deposited
in the Kitakyushu Museum of Natural His-
tory, Kitakyushu (KMNH), and additional
paratypes in the National Museum of Natu-
ral History, Smithsonian Institution, Wash-
ington, D.C. (USNM), and Marine Biologi-
cal Laboratory, Faculty of Fisheries, Kago-
shima University, Kagoshima (KUMB).
Measurements shown in parentheses under
*‘Material examined”’ indicate the maximum
carapace width in millimeters. Abbreviations
used include: M, male; F female.
Family Potamidae Ortmann, 1896
Genus Geothelphusa Stimpson, 1858
Geothelphusa marmorata, new species
Figs. 1-3, Tables 1-3
Material examined.—River Anboh: Ar-
akawa, 1280 m alt., 24 Oct 1998: holotype,
M (29.4), KMNH-IvR 900005, paratype, F
(21.1), KMNH-IvR 900006, M (23.2), F
28.9), USNME 263857 %> 1120 m alt., $ Jul
19975 3) MEG05., 26,6, 21.8), KUMBcr
1053 (used for the electrophoretic analysis).
Diagnosis.—Penultimate segment of
male first gonopod slightly curved laterally,
ultimate segment strongly curved mesially,
cone-shaped, with terminal aperture. Ocular
peduncle swollen proximally, cornea small.
Gastric cristae stout, H-shaped median gas-
tro-cardiac depression distinct. In life, car-
apace and pereopods chocolate brown or
dark red with scattered black speckles.
Description.—Carapace much broader
than long, smooth, devoid of setae (Fig.
la); faint, short oblique striae on epibran-
chial and posterolateral regions; epi- and
uro-gastric regions distinct, former divided
into 2 stout gastric cristae by deep median
groove; H-shaped median gastro-cardiac
depression deep, wide; deep transverse
S2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
view; c, ventral view. Male paratype (USNM 268571): d, ventral view; e, dorsal view. Scales indicate 10 mm.
groove between cardiac and intestinal re-
gions; cervical groove obsolete on epibran-
chial region. Anterolateral margin of cara-
pace cristate, lined with fine rounded gran-
ules; epibranchial notch rudimentary. Pos-
terior margin of epistome divided into 3
parts by 2 deep notches (Fig. 1b), granules
present on lower edge of epistome, absent
medially. Lower orbital margin and groove
between subhepatic and pterygostomian re-
gions lined with faint granules.
Geothelphusa marmorata, new species, male holotype (KMNH-IvR 900005): a, dorsal view; b, frontal
Eyestalk short, proximally swollen, distally
slender. Cornea small, slightly wider than dis-
tal portion of ocular peduncle (Fig. la, b).
Merus of third maxilliped broad, squar-
ish, with deep depression (Fig. 1b). Palp 3-
segmented, connected on inner distal angle
of merus, tip of palp not below distal mar-
gin of ischium. Exopod slender, longer than
ischium, with small 5-segmented flagellum
(Figs. lc, d, 2a; exopodal flagellum dam-
aged in holotype).
VOLUME 113, NUMBER 1
33
Fig. 2.
Geothelphusa marmorata, new species, male paratype (USNM 268571): a, exopod of third maxil-
liped, frontal view; b—g, male holotype (KMNH-IvR 900005): b, left mandibular palp, ventral view; c, left first
gonopod, dorsal view; d, same, ventral view; e, left second gonopod, dorsal view; f, same, ventral view; g. tip
of left second gonopod, dorsal view. Scales indicate | mm.
Chelipeds asymmetrical in males, sym-
metrical in females; movable finger of large
cheliped strongly curved in large male (not
so in small male) (Fig. la, c—e); palm smooth,
outer surfaces convex (Fig. la, c, d). Carpus
of large cheliped almost smooth, with stout
inner tooth, below which is a low projection
(Fig. la, e). Carpus of small cheliped without
any projection below stout inner tooth.
Palp of mandible 3-segmented (Fig. 2b);
distal segment uniramous, sickle-shaped;
median segment longer than wide, distal
34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
half expanded; proximal segment short,
stout.
Penultimate segment of adult male first
gonopod (G 1) gently curved laterally (Fig.
2c, d), synovial membrane short, about 3
times as long as broad (Fig. 2c), ultimate
segment strongly curved mesially (Fig. 2c,
d), cone-shaped, with terminal aperture.
Male second gonopod (G 2) slender, flat,
weakly convex at proximal part, a small
cup-like structure on distal one-fourth, tip
of GZ coneave (Fis: Ze: f.1¢). GZ shorter
than G 1, tip of G 2 not protruding from
aperture of G 1 when coupled.
Color in life-—Carapace and pereopods
(Fig. la, e) chocolate brown or dark red
with scattered black speckles. Lower part of
palm and inmovable finger of both cheli-
peds white in large male, movable finger
and upper part of palm of chelipeds choc-
olate brown. In small males and females,
lower part of palm of both chelipeds white,
fingers and upper part of palm dark red.
Otherwise, no color variation observed be-
tween sexes and sizes.
Genetic characteristics—Among fifteen
gene loci coding for twelve enzymes and
one protein, allelic substitution was ob-
served between G. marmorata, new spe-
cies, and G. exigua at Lactate dehydroge-
nase (LDH*), Isocitrate dehydrogenase-1
(IDHP-1*), and Phosphogluconate dehy-
drogenase (PGDH*) loci (Table 2), and be-
tween G. marmorata and G. dehaani at
PGDH* locus. Nei’s genetic distances (D)
were calculated between all samples based
on the fifteen loci (Table 3). The D values
of G. dehaani populations and G. exigua
populations were low (0.016—0.066 and
0.037, respectively). However, the D values
between G. marmorata and G. dehaani
were high, ranging from 0.155 to 0.2, and
the values between G. marmorata and G.
exigua were higher (0.473 and 0.534).
Etymology.—The specific name is de-
rived from the Latin marmoratus, marbled,
alluding to the characteristic color pattern
of the new species.
Remarks.—The saber-like G 1 and the 3-
segmented mandibular palp with uniramous
distal segment present in the new species
are characteristics of the genus Geothelphu-
sa (see Bott 1970). The medium-sized car-
apace, anterolateral margin lined with small
rounded granules and absence of an epi-
branchial tooth ally G. marmorata with G.
dehaani, G. exigua, G. bicolor, G. miya-
zakii, G. candidiensis, G. ferruginea, G.
tali, G. shokitai, and G. minei, from which
it is distinguished by several features.
The life color of the carapace with scat-
tered black speckles easily distinguishes G.
marmorata from G. dehaani, G. bicolor, G.
miyazakii, G. ferruginea, and G. tali. The
most definitive differences are in the struc-
ture of the G 1. The G 1 ultimate segment
in eight related species, except for G. exi-
gua, is Straight or slightly curved mesially
(Bott 1967, 1970; Minei 1973, 1974a; Su-
zuki & Tsuda 1994, Shy et al. 1994, Shy &
Ng 1998). The ultimate segment in G. exi-
gua is curved laterally and tapering, and
has a subterminal aperture. However, in G.
marmorata, this segment is strongly curved
mesially, ending in a papilla-like tip with a
terminal aperture. The eyestalks in the eight
related species are constricted medially, and
the cornea and proximal part of the ocular
peduncle are swollen. In G. marmorata,
only the proximal part of the ocular pedun-
cle is swollen, while the cornea and the dis-
tal part of the peduncle are proportionately
narrower as in G. exigua.
Previous genetic studies revealed that the
different populations of G. dehaani in Japan
exhibit varying allele frequencies in some
gene loci in Japan (Sugawara & Gamo
1984, Nakajima & Masuda 1985, Aotsuka
et al. 1995, Ikeda et al. 1998). If allelic sub-
stitution is observed at any gene locus
among the different morphological groups
based on coloration, shape of G 1, and so
on, there is a possibility of the groups ac-
tually representing different species. In the
previous studies, there was no allelic sub-
stitution in any gene locus among the dif-
ferent morphological groups. However, Ike-
da et al. (1998) observed an allelic replace-
VOLUME 113, NUMBER 1 35
Table 2.—Allele frequencies at 15 loci for 3 populations of G. dehaani, 2 populations of G. exigua and |
population of G. marmorata in Kagoshima Prefecture.
G. dehaani G. exigua
G. marmorata
Locus Allele Koutsuki Hitotsutani Anboh Hitotsutani Kamiharai Anboh
AAT* he 0.027 0.025 0.000 0.975 1.000 0.059
*b 0.922 0.975 0.929 0.025 0.000 0.941
Fc 0.051 0.000 0.071 0.000 0.000 0.000
AK* ia 0.961 1.000 1.000 1.000 0.750 1.000
al 2 0.028 0.000 0.000 0.000 0.250 0.000
sa 0.011 0.000 0.000 0.000 0.000 0.000
GAPDH* *q 1.000 1.000 1.000 1.000 1.000 1.000
GPI* *q 0.253 0.025 0.000 0.600 0.200 0.059
*b 0.726 0.475 0.000 0.000 0.275 0.647
=C 0.018 0.050 0.000 0.400 D525 0.000
*d 0.003 0.425 1.000 0.000 0.000 0.235
ye 0.000 0.025 0.000 0.000 0.000 0.059
HK* *a 0.085 0.000 0.000 0.000 0.000 0.000
=D 0.755 1.000 0.786 0.917 0.975 0.735
KG 0.160 0.000 0.214 0.083 0.025 0.265
IDHP- 1* *q 0.891 1.000 1.000 0.000 0.000 1.000
*D 0.070 0.000 0.000 0.000 0.000 0.000
at 3 0.039 0.000 0.000 0.000 0.000 0.000
a 7) 0.000 0.000 0.000 1.000 1.000 0.000
IDHP-2* re) 1.000 1.000 1.000 0.050 0.000 1.000
*b 0.000 0.000 0.000 0.950 1.000 0.000
EDH* na 1.000 1.000 1.000 0.000 0.000 1.000
2 0) 0.000 0.000 0.000 1.000 1.000 0.000
MDH-1* *q 0.964 0.900 1.000 0.975 0.950 1.000
*b 0.036 0.100 0.000 0.025 0.050 0.000
MDH-2* a 0.994 0.975 1.000 1.000 1.000 1.000
2) 0.006 0.025 0.000 0.000 0.000 0.000
nC 0.000 0.000 0.000 0.000 0.000 0.000
ME* “a 1.000 1.000 1.000 1.000 1.000 1.000
MPI* ne 1.000 1.000 1.000 1.000 1.000 1.000
PGDH* ta 0.000 0.000 0.000 0.000 0.000 1.000
4 0.991 1.000 1.000 1.000 1.000 0.000
ss 0.009 0.000 0.000 0.000 0.000 0.000
PGM* wa 0.875 0.925 Oey | 0.925 0375 0.971
=D 0.018 0.075 0.214 0.075 0.625 0.000
=G 0.108 0.000 0.214 0.000 0.000 0.029
PROT* a 1.000 1.000 1.000 0.000 0.000 0.059
*D 0.000 0.000 0.000 1.000 1.000 0.941
Table 3.—Genetic distance among 3 populations of G. dehaani, 2 populations of G. exigua and | population
of G. marmorata in Kagoshima Prefecture.
G. dehaani G. exigua
Koutsuki Hitotsutani Anboh Hitotsutani Kamiharai
(1) (II) (IIT) (IV) (V)
G. dehaani (11) 0.016
(IIT) 0.032 0.066
G. exigua (IV) 0.469 0.462 0.519
(V) 0.520 0.518 0.559 0.037
G. marmorata Anboh (VI) 0.156 0.155 0.200 0.473 0.534
36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
130° 30 E
ips, 3:
130° 40 E
30° 20° N
Distribution and abundance of G. marmorata, new species (solid circle) and G. dehaani (White,
1847) (white circle) in Yakushima Island, Kagoshima, Japan. Arabic numerals correspond to size of circles in
the square at bottom left, and indicate the number of crabs captured per ten minutes by one person. Broken line
shows 1000 m contour line.
ment at three loci between color morphs
representing allopatric populations. No
morphological differences in G 1 structure
have been reported among those popula-
tions thus far. In this study, allelic substi-
tution was observed at three loci among G.
marmorata, G. dehaani, and G. exigua. In
addition, G. marmorata, and G. dehaani are
sympatric in Anboh River, Yakushima Is-
land. The observed allelic substitutions,
therefore, strongly indicate the presence of
reproductive isolation among three species.
The genetic distance (D values) further sug-
gest that G. marmorata, belongs to a dif-
ferent evolutionary lineage from the Anboh
population of G. dehaani.
Distribution.—The specimens of G. mar-
morata, examined have been obtained only
in the area above 950 m altitude on Yaku-
shima Island, Kagoshima Prefecture (Fig.
3). Geothelphusa marmorata, and G. de-
haani are sympatric, having been taken to-
VOLUME 113, NUMBER 1
gether at some locations from 950 m to
1350 m altitude.
Acknowledgments
We thank C. L. McLay of the University
of Canterbury, and P. K. L. Ng of National
University of Singapore for their critical
reading of the manuscript. Thanks are also
extended to R. Lemaitre for his valuable
comments on the manuscript. We are also
indebted to Y. Hiwatashi for his technical
help with the electrophoretic analysis.
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. 1974a. Potamoid crabs of Taiwan, with de-
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. 1974b. Studies on the freshwater crabs of
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38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):39-—47. 2000.
A new species of the genus Neostylodactylus Hayashi & Miyake, 1968
(Crustacea: Decapoda: Stylodactylidae) from southern Japan
Junji Okuno and Hiroyuki Tachikawa
Coastal Branch of Natural History Museum and Institute, Chiba, 123 Yoshio, Katsuura, Chiba
299-5242, Japan
Abstract.—A new species of stylodactylid shrimp, Neostylodactylus litoralis,
is described and illustrated on the basis of four ovigerous females collected from
sublittoral zones on the Ogasawara and Ryukyu Islands at depths of 3—10.5 m.
This new species is readily distinguished from the five described species of
Neostylodactylus Hayashi & Miyake in having a non-produced posterior margin
of the telson, the noticeably elongate mesialmost pair of posterior spines on the
posterior margin of the telson, and unarmed meri of the third to fifth pereiopods.
Members of the caridean shrimp family
Stylodactylidae are classified in five genera
(Hanamura & Takeda 1996), with species
known to occur in deep waters exceeding
100 m (Chace 1983; Cleva 1990, 1994,
1997). Only one stylodactylid, Neostylodac-
tylus amarynthis (De Man, 1902), has been
recorded in shallow waters less than 100 m
(Kemp 1925, Chace 1983, Cleva 1990).
While sampling the sublittoral zone (3.0—
10.5 m) of the Ogasawara and Ryukyu Is-
lands, southern Japan, we obtained four sty-
lodactylid specimens referable to the genus
Neostylodactylus Hayashi & Miyake, 1968.
Morphologically, our specimens differed
from the five previously described species
of Neostylodactylus, and clearly represent a
new species described here.
Specimens were collected by using a hand
net, and are deposited in the Coastal Branch
of Natural History Museum and _ Institute,
Chiba (CMNH). The method of measure-
ments follows Cleva (1990). The abbrevia-
tion CL indicates postorbital carapace length.
Family Stylodactylidae
Genus Neostylodactylus Hayashi &
Miyake, 1968
Neostylodactylus litoralis, new species
Figs. 1—4
Type series.—Holotype: ovig. 2 CL
29mm (CMNH-Z2C 00071), 27°11.5'N,
142°07.0'E, Takinoura, Ani-jima Island,
Ogasawara Islands, 5 m, Aug 1996, coll. Y.
Morita. Paratypes: 1 ovig. 2 CL 2.8 mm
(CMNH-ZC 00103), 27°04.6'N, 142°07.1’E,
Hyotan-jima Islet, NW of Chichi-jima Is-
land, Ogasawara Islands, 8 m, May 1996,
coll. -T. Gomi. 1 ovig 2 CL 24 mm
(CMNH-ZC 00119), 26°13.7'N, 127°27.4’E,
Gahi-jima Islet, Kerama Group, Ryukyu Is-
lands, 3m, 3, Sep: 1998, coll A. Ono.
Non-type material.—|1 2 CL~2.2 mm
(molting) (CMNH-ZC 00137), 26°42.0’N,
127°27.4'E, Ie-shima Island, Ryukyu Is-
lands, 10.5 m,,14 Jun 1996, coll T. No-
mura.
Diagnosis.—Small sized stylodactylid
species (known specimens CL 2.4-2.9
mm). Carapace armed with supraorbital
spine. Rostrum well developed, falling
short of distal end of scaphocerite, armed
with movable spines on both dorsal and
ventral margins. Pleuron of third to fifth ab-
dominal somite each armed with postero-
lateral spine. Telson armed with 2 pairs of
dorsal and dorsolateral spines, posterior
margin ending roundly, armed usually with
3 pairs of spines, mesialmost pair longest,
with 4 plumose setae between spines. Sca-
phocerite armed laterally with 5 or 6 mov-
able spines. Third to fifth pereiopods with
40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1. Neostylodactylus litoralis, new species. Holotype (CMNH-ZC 00071). A, left carapace with cephalic
appendages, in lateral view. B, right first to sixth abdominal somites, in lateral view. C, telson and uropods, in
dorsal view. D, tip of telson, in dorsal view. E, right antennular peduncle, in ventral view. FE right antenna, in
dorsal view. Scales equal 1.5 mm (A, B), 1 mm (C, E, F), 0.5 mm (D).
VOLUME 113, NUMBER 1 4]
<
ari
NN wat
S
VALLE
\
pe,
ig
Z
C D —
eS
= Se ae
| Ty
fy,
qfeaBi® ifs, / E
F ~
a ee ae: >
Fig. 2. Neostylodactylus litoralis, new species. Holotype (CMNH-ZC 00071). A, right mandible. B, right
maxillule. C, right maxilla. D, right first maxilliped. E, right second maxilliped. E right third maxilliped. Scales
equal 0.5 mm (A, B, C, D), 1 mm (E, F).
uniunguiculate dactyli, armed posteriorly postorbital region slightly concave; infra-
with 3 movable spines; meri unarmed. orbital margin anteriorly produced in tri-
Description.—Carapace (Fig. 1A) _ angular process, distinctly overreaching tip
smooth, glabrous; supraorbital spine acute, of antennal spine; hepatic depression indis-
continuous with feeble postorbital ridge; tinct; antennal spine submarginal, directed
42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1.—Branchial formula of Neostylodactylus
litoralis, new species (female only).
Maxillipeds
I II Ill I II OE IN NY
Pereiopods
Pleurobranchs —_—- — — ] ] ] l ]
Arthrobranchs —
Podobranchs -—
Epipods |
Exopods ]
]
|. . 3 a eae
|
anteriorly; pterygostomian angle armed
with spine. Rostrum (Fig. 1A) straight,
slightly descending, 0.71—0.79 times as
long as carapace; dorsal margin armed with
6—12 articulated teeth, 3—9 teeth anterior to
level of postorbital margin, more or less
equidistant, interspace with simple long
plumose setae, 1 tooth just above level of
postorbital margin; dorsal carina with 2
teeth posterior to level of postorbital mar-
gin, and with 2 or 3 robust plumose setae;
ventral margin armed with 1-3 articulated
teeth.
Abdominal somites (Fig. 1B) smooth,
glabrous; first somite with pleuron sparsely
fringed with simple setae; pleuron of sec-
ond somite fringed with robust plumose se-
tae along anterolateral and ventromesial
margins, with simple setae ventrally; pleu-
ron of third somite armed with small acute
tooth posterolaterally, densely fringed with
robust plumose setae ventrally; pleuron of
fourth somite armed with acute tooth pos-
terolaterally, sparsely fringed with robust
plumose setae marginally; pleuron of fifth
somite armed with acute tooth posterolat-
erally, with plus 2 or 3 elongate stout setae
bearing setules, and simple setae ventrally;
sixth somites 0.33—0.46 times as long as
carapace, unarmed posteroventrally.
Telson (Fig. 1C) 0.52—0.68 times as long
as carapace, dorsal surface armed with 2
pairs of spines, posterior pair situated more
laterally than anterior pair, midline with
transverse row of long plumose setae prox-
imally; posterior margin (Fig. 1D) feebly
rounded, not produced in acute median pro-
cess, armed with 3 pairs of spines (an extra
spine on left side in holotype), mesialmost
pair noticeably elongate, flanking 4 stout
plumose setae.
Eye with lightly pigmented cornea, and
plumose setae, without ocellus; corneal di-
ameter 0.21—0.29 times as long as carapace;
stalk slender.
Antennular peduncle (Fig. 1E) slender,
slightly overreaching or falling slightly
short of rostral apex; proximal segment
armed with ventromesial spine and long
spiniform seta posterior to spine, dorsodis-
tal margin fringed with short simple setae;
stylocerite reaching midlength of proximal
segment, tapering distally in acute point; in-
termediate segment armed dorsolaterally
with 2 spiniform setae; distal segment short,
about half length of intermediate segment.
Upper flagellum ventrally with short setae,
seventh article with long spiniform seta dis-
tolaterally; seventh article of lower flagel-
lum with long, plumose seta.
Antenna (Fig. 1F) with scaphocerite with
lateral margin distinctly concave, slightly
overreaching rostral apex, 0.69—0.75 times
as long as carapace, lateral margin armed
with 5 or 6 acute movable spines, distolat-
eral tooth distinctly overreaching rounded
distal blade; carpocerite fringed with long
simple setae distomesially; basicerite armed
with spine distolaterally; antennal flagellum
armed with long spiniform setae, articula-
tions much indistinct.
Mandible (Fig. 2A) without palp; incisor
process well developed, slightly rounded,
distal margin armed with 5 blunt teeth; mo-
lar process truncated distally, with short
Sparse setae.
Maxillule (Fig. 2B) with feebly bilobed
palp, inner lobe with long simple seta,
armed dorsally with small spine proximal
to outer lobe; upper lacinia fringed with nu-
merous setae, distal 3 setae considerably
longer, plumose; lower lacinia distally with
numerous simple setae, midlength of mesial
margin with single short seta.
Maxilla (Fig. 2C) with palp distally
fringed with long plumose setae; basal en-
dite bilobed, with numerous setae on mesial
VOLUME 113, NUMBER 1 43
tc ——
>
Spe)
= hia
LL
i)
(é
)
6
TY
/
5M
ii eA
=
ai atin eras 5)
Fig. 3. Neostylodactylus litoralis, new species. Holotype (CMNH-ZC 00071). A, left first pereiopod. B, left
second pereiopod. C, left third pereiopod. D, same, dactylus. Scales equal 1.5 mm (A, B), | mm (C), 0.5 mm
(D).
44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Neostylodactylus litoralis, new species. A, holotype (CMNH-ZC 00071), fresh specimen, lateral view
(photo by H. Tachikawa); B, paratype (CMNH-ZC 00119), alive in aquarium (photo by A. Ono).
margin, upper lobe slightly overreaching First maxilliped (Fig. 2D) with well de-
level of distal margin of lower lobe; coxal veloped exopod; caridean lobe well devel-
endite feebly rounded, with numerous setae. oped, rounded, palp slender, setose; basal
Scaphognathite broad, rounded, marginally endite distally truncate, with 3 rows of nu-
with numerous setae. merous setae; coxal endite distinct, with 2
VOLUME 113, NUMBER 1
rows of numerous setae mesially. Epipod
bilobed.
Second maxilliped (Fig. 2E) with well
developed exopod; terminal 2 segments im-
planted side by side at distal end of ante-
penultimate segment, flexor segment slight-
ly longer and narrower than extensor seg-
ment, distally fringed with numerous sim-
ple setae, flexor margin with 3 simple setae,
extensor segment rounded, marginally with
numerous long simple setae; antepenulti-
mate segment protruded dorsodistally, with
long simple setae, external and mesial mar-
gins with long simple setae; carpal segment
short, naked; ischiomeral segment with dor-
sal margin sparsely with setae with setules,
mesially with similar setae. Epipod oblong,
with well developed podobranch.
Third maxilliped (Fig. 2F) slender, over-
reaching distal margin of scaphocerite by
full length of ultimate segment, without ex-
opod; flexor margins of distal 3 segments
densely fringed with long setae with nu-
merous setules and sparsely with short plu-
mose setae; penultimate segment |.21—1.22
times as long as ultimate segment, armed
with row of 5-8 articulated spines; ante-
penultimate segment with lateral row of 9-
12 articulated spines, and | spine on dorsal
surface. Small arthrobranch present.
Branchial formula as indicated in Table 1.
First pereiopod (Fig. 3A) overreaching
distal end of scaphocerite by distal margin
of carpus; chela with reduced palm about
one-fifth of movable and fixed fingers,
proximally convex, cutting edges of both
fingers entire, without tooth, movable finger
with flexor margin fringed with long setae
with numerous setules, fixed finger fringed
ventrally with similar setae; carpus 0.83-—
0.93 times as long as carapace, dorsal mar-
gin armed with row of spiniform setae, lat-
eral surface armed with row of spines, ven-
tral margin fringed with long setae with nu-
merous setules and with a few plumose
setae; ischiomerus armed laterally with row
of spines, with short simple setae distola-
terally, ventral margin fringed with long se-
45
tae with numerous setules and with a few
plumose setae.
Second pereiopod (Fig. 3B) overreaching
distal end of scaphocerite by proximal mar-
gin of movable finger; chela with reduced
palm about 0.20 times as long as fingers,
proximally convex, cutting edges of both
fingers entire, without tooth, movable finger
with flexor margin fringed with long setae
with numerous setules, fixed finger fringed
ventrally with similar setae; carpus 0.67—
0.79 times as long as carapace, dorsal mar-
gin armed with row of spiniform setae, lat-
eral surface armed with row of spines, ven-
tral margin densely fringed with long setae
with numerous setules and with a few plu-
mose setae; ischiomerus armed laterally
with row of spiniform setae, distolaterally
with spine and short plumose setae, ventral
margin densely fringed with long setae with
numerous setules and sparsely with long
plumose setae.
Third pereiopod (Fig. 3C) slightly over-
reaching midlength of scaphocerite; dacty-
lus (Fig. 3D) uniunguiculate, armed with 3
movable spines posteriorly; propodus 0.75—
0.76 times as long as carapace, 2.20—2.25
times as long as carpus, distomesial margin
concave, ventrally armed with short spines;
carpus unarmed, with robust plumose setae
distolaterally; ischiomerus unarmed, 0.76—
0.83 times as long as carapace, 2.20—2.50
times as long as carpus, laterally with long
robust plumose setae, dorsodistal margin
with short plumose setae, ventrally with
short plumose setae. Fourth and fifth pe-
reiopods similar to third pereiopod. Fourth
pereiopod falling slightly short of mid-
length of scaphocerite. Fifth pereiopod
overreaching basicerite by full length of
dactylus.
Uropod (Fig. 1C) slightly overreaching
distal margin of telson. Protopodite armed
with strong lateral spine; exopod armed
with acute fixed tooth distolaterally, mesi-
ally with stout movable tooth; endopod
with prominent lateral lobe proximally.
Color in life (Fig. 4).—Carapace and ab-
dominal somites reddish-violet, posterolat-
46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
eral part of carapace and dorsolateral parts
of the first to fourth somites darker than
other parts, covered with numerous fine red
spots. Antennular peduncle transparent-red-
dish, flagella transparent-whitish. Scapho-
cerite transparent, antennal basicerite red-
dish-violet. Telson, third maxilliped, ante-
rior two pereipopods and uropods transpar-
ent. Ambulatory pereiopods with coxae and
basis reddish-violet with numerous red
spots; meri, carpi, propodi and dactyli
transparent. Pleopods transparent.
Etymology.—litoralis, from Latin Jitus,
the shore, in allusion to the habitat in a re-
markably shallow depth in the vertical dis-
tributional range of Stylodactylidae.
Distribution.—Known only from the
Ogasawara and Ryukyu Islands, southern
Japan.
Remarks.—Based on the distally acute
stylocerite, the prominent supraorbital
spine, and the absence of the mandibular
palp and arthrobranchs on all pereiopods,
the specimens clearly belong in the genus
Neostylodactylus.
The rostral armature does show marked
intraspecific variations. The holotype is
armed dorsally with six teeth anterior to the
level of the postorbital margin. The dorsal
margin anterior to the orbital margin has
nine teeth in the paratype from the Ogasa-
wara Islands (CMNH-ZC 00103), and three
teeth in that from the Ryukyu Islands
(CMNH-ZC 00119).
The non-type material (CMNH-ZC
00137) was molting when captured.
Collectors of the present specimens ob-
served that WN. litoralis appeared to face up-
wards. The carapace was bent strongly
backwards towards the abdominal somites,
with the dorsal surface of the carapace al-
most in contact with the somites (see Fig.
4B).
In recent taxonomic studies dealing with
this genus (Chace 1983; Cleva 1990, 1994,
1997; Komai 1997), five Neostylodactylus
species were recognized: N. affinis Hayashi
& Miyake, 1968; N. amarynthis (De Man,
1902); N. hayashii Komai, 1997; N. inves-
tigatoris (Kemp, 1925); and N. sibogae (De
Man, 1918).
The non-produced posterior margin of
the telson and the unarmed meri of the am-
bulatory pereiopods readily distinguish WN.
litoralis from all other congenerics. In other
congeneric species, the posterior margin of
the telson is acutely produced mesially, and
the meri of ambulatory pereiopods are each
armed with one or more lateral spines (see
De Man 1918, 1920; Kemp 1925, Hayashi
& Miyake 1968, Chace 1983, Komai 1997).
Moreover, Dr. R. Cleva (in litt.) kindly in-
formed us that the long mesialmost pair of
spines on the posterior margin of the telson
is also a characteristic of this new species.
Furthermore, N. litoralis differs from N.
amarynthis, the type species of the genus,
and N. affinis, in having the rostrum falling
short of the distal end of the scaphocerite,
and the telson armed with two pairs of
spines instead of five and three respectively.
The smooth pleura of the first and second
abdominal somites and the lack of the spine
on the lateral surface of the sixth abdominal
somite also separate the present new species
from N. amarynthis. The rostral formula
separate N. litoralis from N. affinis. From
N. investigatoris and N. sibogae, the present
new species is readily distinguished by hav-
ing an armed ventral margin of the rostrum,
and the presence of marginal spines on the
third to fifth abdominal somites. Neostylo-
dactylus litoralis differs from N. hayashii
by having a much shorter rostrum, which
falls short of the distal end of the scapho-
cerite; the rostral armature; the lateral mar-
gin of the scaphocerite armed with spines;
and the number of spines on the dorsal sur-
face of the telson.
Acknowledgements
We thank the Japanese skillful divers,
Messrs. T. Gomi, Y. Morita, T. Nomura, and
A. Ono, for making the material available
for this study. Mr. A. Ono kindly permitted
us to use his color photograph. Drs. A. J.
Bruce and R. Cleva kindly read an early
VOLUME 113, NUMBER 1
draft, and provided valuable comments. The
manuscript was benefited from reviews by
Drs. T. Komai and R. Lemaitre.
Literature Cited
Chace, FE A., Jr. 1983. The caridean shrimps (Crusta-
cea: Decapoda) of the Albatross Philippine Ex-
pedition, 1907-1910, part 1: Family Stylodac-
tylidae.—Smithsonian Contributions to Zoolo-
gy 381:1-21.
Cleva, R. 1990. Crustacea Decapoda: Les genres et les
espéces indo-ouest pacifiques de Stylodactyli-
dae. Pp. 71-136 in A. Crosnier, ed., Résultats
des Campagnes MUSORSTOM, 6.—Mémoires
du Muséum National d’ Histoire Naturelle, Paris
(A) 145.
. 1994. Some Australian Stylodactylidae (Crus-
tacea: Decapoda), with descriptions of two new
species.—The Beagle, Records of the Museums
and Art Galleries of the Northern Territory 11:
53-64.
. 1997. Crustacea Decapoda: Stylodactylidae
récoltés en Indonésie, aux iles Wallis et Futuna
et au Vanuatu (Campagnes KARUBAR, MU-
SORSTOM 7 et 8). Données complémentaires
sur les Stylodactylidae de Nouvelle-Calédonie.
Pp. 385—407 in A. Crosnier & P. Bouchet, eds.,
Résultats des Campagnes MUSORSTOM,
16.—Mémoires du Muséum National d’ Histoire
Naturelle, Paris 172.
De Man, J. G. 1902. Die von Herrn Professor Kiiken-
thal im Indischen Archipel gesammelten De-
47
kapoden und Stomatopoden. /n W. Kiikenthal,
Ergebnisse einer zoologischen Forschungsreise
in den Molukken und Borneo.—Abhandlungen
Herausgegeben von der Senckenbergischen Na-
turforschenden Gesellschaft 25:467—929 pls.
19-27.
. 1918. Diagnoses of new species of macrurous
decapod crustacea from the Siboga-Expedi-
tion.—Zoologische Mededelingen, Leiden 4:
159-166.
. 1920. The Decapoda of the Siboga Expedi-
tion, part IV. Families Pasiphaeidae, Stylodac-
tylidae, Hoplophoridae, Nematocarcinidae,
Thalassocaridae, Pandalidae, Psalidopodidae,
Gnathophyllidae, Processidae, Glyphocrangon-
idae, and Crangonidae.—Siboga-Expeditie
39a3:1-318, pls. 1-25.
Hanamura, Y., & M. Takeda. 1996. Establishment of a
new genus Bathystylodactylus (Crustacea: De-
capoda: Stylodactylidae), with description of a
new species from northwestern Pacific.—Zoo-
logical Science 13:929—934.
Hayashi, K.-I., & S. Miyake. 1968. Notes on the fam-
ily Stylodactylidae with the description of a
new genus Neostylodactylus.—Journal of the
Faculty of Agriculture, Kyushu University 14:
583-611.
Kemp, S. 1925. Notes on Crustacea Decapoda in the
Indian Museum, XVII: On various Caridea.—
Records of the Indian Museum 27:249-—343.
Komai, T. 1997. A new species of the shrimp genus
Neostylodactylus Hayashi and Miyake (Crusta-
cea: Decapoda: Stylodactylidae) from Japan.—
Natural History Research 4:125—133.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):48-53. 2000.
On the male of Scutumara enodis Ng & Nakasone, 1993
(Crustacea: Decapoda: Brachyura: Grapsidae)
N. K. Ng and T. Komai
(NKN) Department of Biological Sciences, National University of Singapore, Lower Kent Ridge
Road, Singapore 119260, Republic of Singapore; (TK) Natural History Museum & Institute
Chiba, 955-2 Aoba-cho, Chuo-ku, Chiba 260, Japan
Abstract.—The male of the unusual grapsid, Scutumara enodis Ng & Na-
kasone, is described for the first time, and the species is compared with its
congeners, S. laniger (Tesch) and S. miyakei (Nakamura & Takeda). The male
characters of S. enodis, the type species of the genus, provide further evidence
to support the establishment of Scutumara Ng & Nakasone, for these three
species. The male abdomen and male first gonopods of S. enodis are, however,
distinctly different from those of S. laniger and S. miyakei.
A male specimen of the coral reef grap-
sid crab, Scutumara enodis Ng & Naka-
sone, 1993, was recently collected by the
second author in Ishigaki Island, Yaeyama
Group, the Ryukyus. This species was orig-
inally described from only a single subadult
female. This study describes the male spec-
imen of this species, and gives information
on the male abdomen and male first pleo-
pod (G1). Our observations provide addi-
tional evidence in support of the establish-
ment of the genus Scutumara Ng & Naka-
sone, 1993.
Measurements are given in mm in the or-
der: carapace width by length. Specimens
used here are deposited in the Zoological
Reference Collection (ZRC), Department of
Biological Sciences, National University of
Singapore; Natural History Museum & In-
stitute (CBM), Chiba, Japan; and National
Museum of Natural History, Smithsonian
Institution, Washington, D.C., U.S.A.
(USNM).
Family Grapsidae MacLeay, 1838
Subfamily Varuninae Alcock, 1900
Scutumara enodis Ng & Nakasone, 1993
(Figse2d32)
Scutumara enodis Ng & Nakasone, 1993:
1, figs. 1-2.
Material examined.—Holotype, female
(5.8 X 5.9 mm) (ZRC.1993.1), under litto-
ral coral sand, Kunri-Hama Beach, Sesoko
Island, Okinawa, Ryukyus, Japan, coll.
PK.L. Ng & Y. Nakasone, Apr 1992. Oth-
ers—l1 male (4.9 X 5.1 mm) (CBM-ZC
3604), beach near Fukido-gawa river-
mouth, Ishigaki Island, Yaeyama group, Ja-
pan, coll. T. Komai, 24 Mar 1997.
Description of male.—Carapace slightly
longer than broad; dorsal surface smooth,
glabrous, without setae; regions not defined,
strongly convex transversely and longitu-
dinally, gastric region most convex (Fig.
1A). Frontal margin slightly convex, entire;
lateral lobes not visible dorsally, slightly
deflexed downwards; lateral edges appar-
ently confluent with supraorbital margin
from dorsal view, but not confluent from
frontal view; inner edges of smooth, entire
supraorbital margins strongly deflexed
downwards, much more than outer edges of
front, forming slight crimp at junction be-
tween frontal and supraorbital margins. In-
fraorbital margin not distinctly cristate,
slightly raised; not granulated or striated.
Anterolateral margin slightly arcuate, sub-
cristate, very faintly trilobed, lobes separat-
ed by very broad, shallow clefts; external
orbital lobe most distinct, very broad; sec-
VOLUME 113, NUMBER 1
ond, third lobes more like weak undula-
tions. Posterolateral margins not sharply de-
marcated from anterolateral margin, ap-
pearing almost straight, distinctly converg-
ing. Orbits small, eyes completely filling
orbit, corneae well developed. Pterygosto-
mial, suborbital, branchial regions smooth.
Orbital hiatus completely filled by large
basal anntennal segment; flagellum com-
pletely enclosed within orbit. Antennules
folding obliquely, fossae very large; basal
segment large, subtriangular in shape,
broader than long. Posterior margin of ep-
istome weakly sinuous, with 3 distinct ridg-
es (1 median, 2 lateral); ridges separated
from each other by narrow gap. Endosto-
mial region with 3 well developed longi-
tudinal palatal ridges (1 median, 2 lateral);
lateral palatal ridges joining inner edges of
lateral ridges of posterior epistomial mar-
gin.
Third maxilliped with foliaceous merus,
broader than long; anterolateral angle
strongly produced, auriculiform; distal mar-
gin distinctly bilobed, outer lobe larger, its
base with small median cleft. Ischium lon-
ger than broad, sulcus not discernible.
Small, distinct rhomboidal gape formed be-
tween inner margins of meri and ischia of
third maxillipeds when closed. Exopod with
obtuse, blunt inner subdistal angle, flagel-
lum longer than width of merus (Fig. 1B).
Chelipeds small, subequal, outer, inner
surfaces smooth, glabrous; merus, carpus
without spines or teeth; inner distal angle
of carpus with broad, low, rounded lobe.
Outer surface of chela with low but distinct
ventral ridge running from near proximal
part of palm to almost tip of pollex (Fig.
1C); inner surface without setae at base of
fingers. Fingers distinctly longer than palm;
cutting edge of both fingers with numerous
denticles, ending in recurved, sharp tips; no
gape discernible when fingers closed.
Ambulatory legs with second pair lon-
gest (Fig. 1D). All segments smooth, with-
out spines or setae. Dorsal and ventral mar-
gins of merus sub-cristate, dorsal margin
49
with blunt subdistal tooth. Dactylus taper-
ing to slender, acute tip (Fig. 1E).
Male abdomen triangular (Fig. 1F); lat-
eral margins sparsely setose; first abdomi-
nal segment weakly arched, with very weak
transverse ridge; second segment narrow,
short; third segment broad, slightly swollen
laterally but medially depressed with prox-
imal margin broader than distal margin, lat-
eral margins rounded; fourth segment
broader but shorter than fifth segment; fifth
segment with proximal and distal margins
straight, lateral margins slightly concave;
sixth segment quadrate, lateral margins
trapezoidal, weakly convex, proximal mar-
gin slightly concave, convex medially. Tel-
son sub-triangular, lateral margins weakly
concave, distal margin rounded.
Lateral margins of first 2 thoracic ster-
nites finely granulated; suture between ster-
nites 2 and 3 slightly convex towards ab-
domen; lateral margins of sternites 3 and 4
sinuous, with deep, broad notch demarcat-
ing edge of suture; median groove between
sternites 5 and 6 narrow; space between
Ssternites 8 very narrow throughout length
(Fig. 2A).
Penis located at base of eighth sternite
(Fig. 2A). G1 relatively slender, weakly
curving outwards, reaching to anterior mar-
gin of fifth sternite (Fig. 2B); terminal lobe
elongate, apparently 2-articulated, dorsally
curved; genital opening lateral to base of
terminal lobe; subterminal lobe chitinous,
rounded, narrower than terminal lobe,
densely setose (Figs. C—E). G2 short, small.
Distribution.—Southern Japan (Okinawa
south to Ishigaki Islands, Ryukyus).
Remarks.—Ng & Nakasone (1993) es-
tablished Scutumara and transferred two
species previously placed in Pseudograp-
sus, S. laniger (Tesch, 1918), and S. miyakei
(Nakamura & Takeda, 1972), to this new
genus. Although the type specimen of S.
enodis was represented only by a subadult
female, Ng & Nakasone (1993) argued that
on the basis of differences on the carapace
and ambulatory legs, S. enodis, S. laniger
and S. miyakei are distinct enough to war-
50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
E
Fig. 1. Scutumara enodis Ng & Nakasone, 1993 (male, 4.9 by 5.1 mm, CBM-ZC 3604). A, carapace; B,
third maxilliped; C, dorsal view of left chela; D, second ambulatory leg; E, fourth ambulatory leg; K abdomen.
Scales: A, C = 1.0 mm; B, E, F = 0.5 mm.
VOLUME 113, NUMBER | 5]
Fig. 2. Scutumara enodis Ng & Nakasone, 1993 (male, 4.9 by 5.1 mm, CMB-ZC 3604). A, sternum; B, natural
position of male first gonopod; C, male first gonopod in different view; D, enlarge view of the G1; E, different
views of the shaven enlarged distal region of Gl. Scales: A, B = 1.0 mm, C = 0.5 mm, D, E = 0.25 mm.
D2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
rant their placement in Scutumara. The dis-
covery of the male S. enodis provide further
evidence to the establishment of Scutu-
mara. We have also found that the exopod
of the third maxillipeds of all three species
reaches two-thirds the length of the merus,
while the exopod of the third maxillipeds
in Pseudograpsus species reach to half the
length of the merus, a character that further
seperates these two genera. In addition, all
known Scutumara species are ivory-cream
white when alive, while all Pseudograpsus
species are chestnut in color. The lateral
margins of the sixth abdominal segment are
generally rounded in Scutumara, however,
in Pseudograpsus, the lateral margins are
angular at the distal region. The sixth ab-
dominal segment in Scutumara (length to
width ratio between 3.5 and 5.0) is narrow
compared to Pseudograpsus (length to
width ratio between 1.5 to 2.0). Noteworthy
is that the dentation on the anterolateral
margin of the male specimen of S. enodis
is less prominent than that of the female
holotype. We interpret this as intraspecific
variation.
Scutumara enodis is separated from S.
laniger and S. miyakei by the different
structure of the frontal margin, the absence
of gastric and cardiac grooves, and absence
of setae on the inner surface of the palm
(Ng & Nakasone 1993). Based on pub-
lished descriptions and figures of S. laniger
and S$. miyakei, the male abdomen and gon-
opod of S. enodis are also different. The
third abdominal segment is narrower in S.
enodis (length to width ratio: ca. 4.1) than
for its congeners (ca. 3.5 in S. laniger, ca.
3.1 in S. miyakei). In the two species of
Pseudograpsus examined for this character,
the length to width ratio of the third abdom-
inal segment ranges from ca. 3.8 (P. elon-
gata) to ca. 4.0 (P. albus), which is narrow-
er than S. laniger and S. miyakei, but only
slightly broader than S. enodis. In S. enodis,
the lateral margins of the sixth abdominal
segment are weakly rounded, while in S.
laniger, the posterior part of the lateral mar-
gins are distinctly convex; in S. miyakei the
lateral margins are even more convex. The
sixth abdominal segment of S. enodis is
narrower (length to width ratio: ca. 5.0)
than that of S. laniger (ca. 3.5) and S. mi-
yakei (ca. 3.8). The length to width ratio of
the telson in S. enodis is 1.0, which is sim-
ilar to S. /aniger, but is relatively longer in
S. miyakei (ca. 1.4). Nakamura & Takeda
(1972) commented that the G1 of S. miyak-
ei differs from S. laniger and Pseudograp-
sus elongata (A. Milne-Edwards, 1873) in
having a short distal part and obtuse sub-
terminal lobe, while the latter two have
long beaks and small subterminal lobes
(Nakamura & Takeda 1972: 438). This im-
plies that the G1 of S. laniger is also longer
and more slender than in S. enodis and S.
miyakei. The G1 of S. enodis is character-
ised by having an elongated tube, and nar-
row subterminal lobe. The length of the dis-
tal part of the G1 of Scutumara seem to be
variable within the genus. Likewise, in the
three species of Pseudograpsus examined
(P. elongata, P. albus and P. crassus) for
this character, the length of the distal part
of the G1 varies greatly, from long (P. elon-
gata) to short (P. albus), indicating that the
G1 is useful character at the species level
but not at the generic level.
Specimens of Scutumara enodis are
small, as are the other two known Scutu-
mara species. The holotype female (a sub-
adult) is only 5.8 by 5.9 mm (Ng & Na-
kasone 1993). The smaller adult male spec-
imen, is only 4.9 by 5.1 mm. Similarly, the
holotype of S. laniger is only 8.3 mm in
carapace length, and that of S. miyakei is
3.6 mm. On the other hand, Pseudograpsus
species are generally larger but it is not con-
sistent. The two smallest species are P.
elongata and P. albus: P. elongata ranges
from 8.8 by 7.8 mm (USNM 33411) to 9.7
by 8.6 mm (Crosnier 1965); and P. albus
ranges from 8.2 by 7.3 mm (USNM 81732)
to 9.5 by 8.6 mm (Crosnier 1965). P. cras-
sus (42.6 by 36.3 mm) is a very large spe-
cies (USNM 93152).
VOLUME 113, NUMBER 1
Acknowledgments
The authors are most grateful to Dr. Peter
K. L. Ng and Mr. Yixiong Cai for help in
checking the manuscript. Thanks are due to
Dr. Hiroshi Suzuki (Kagoshima University)
kindly obtained some key references for the
first author, and Mrs. C. M. Yang (ZRC) for
the loan of the holotype. Thanks are due to
the Associate Editor and the two referees
for laboriously correcting the manuscript.
This is contribution 34/98 from the Ecology
and Systematics Laboratory, Department of
Biological Sciences, National University of
Singapore. Support from research grant RP
950324 to Dr. Peter K. L. Ng from the Na-
tional University of Singapore is acknowl-
edged.
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Tesch, J. J. 1918. The Decapoda Brachyura of the Si-
boga Expedition. I. Hymenosomidae, Retro-
plumidae, Ocypodidae, Grapsidae and Gecar-
cinidae.—Siboga-Expeditie 39c, 2:1—148, pls.
1-6.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):54-—65. 2000.
Larval development of Cryptolithodes expansus Miers
(Decapoda: Anomura: Lithodidae) reared in the laboratory
Mi Hyang Kim and Sung Yun Hong
Department of Marine Biology, Pukyong National University, Pusan 608-737, Korea
E-mail: syhong @ dolphin.pknu.ac.kr
Abstract.—The complete larval development of Cryptolithodes expansus
Miers is described based on laboratory rearing. The species has four zoeal
stages and a megalopa. The larvae are described and illustrated, and detailed
comparisons are made with other lithodid larvae, particularly those of Cryp-
tolithodes typicus Brandt.
Cryptolithodes expansus Miers, 1879 is
found along the Korean coast (personal ob-
servation) and in Japan (Makarov 1938,
1962) on rocky bottoms in the sublittoral
region to a depth of about 50—60 m. The
family Lithodidae is represented by 16 gen-
era and 95 species (Dawson 1989), but the
larval development of only 16 species, rep-
resenting 9 genera, has been described
(Konishi 1986; Haynes 1984, 1993; Koni-
shi & Taishaku 1994).
Sars (1890) described all zoeal stages up
to the megalopa of Lithodes maja Linnaeus,
1758. Subsequent studies of Lithodidae lar-
vae have been described, at least in part, for
15 species (Konishi 1986; Haynes 1984,
1993; Konishi & Taishaku 1994): Crypto-
lithodes typicus Brandt, 1848, Dermaturus
mandtii Brandt, 1850, Hapalogaster greb-
nitzkii Schalfeew, 1892, H. dentata (De
Haan 1844), H. mertensii Brandt, 1850,
Lithodes aequispina Benedict, 1895, L. an-
tarctica Jacquinot, 1853, Lopholithodes
manatii Brandt, 1848, Paralithodes brevi-
pes (H. Milne Edwards & Lucas, 1841), P.
camtschaticus (Tilesius 1815), P. platypus
(Brandt 1850), Paralomis granulosa (Jac-
quinot 1852), P. hystrix (De Haan 1846),
Placetron wosnessenskii Schalfeew, 1892,
and Rhinolithodes wonessenskii Brandt,
1848.
Hart (1965) described all zoeal stages,
megalopa and crab 1 stage of Cryptolitho-
des typicus Brandt, 1848 reared in the lab-
oratory. This is the only species of Cryp-
tolithodes for which larvae are known so
far. The purpose of the present study is to
describe complete larval stages of C. ex-
pansus reared in the laboratory, and to com-
pare them with those of C. typicus.
Materials and Methods
On 20 August 1995 a local fisherman
caught three berried females of Cryptolitho-
des expansus with a commercial octopus
trap in the vicinity of Pusan (35°10'N,
129°10’E), Korea. The ovigerous females
were brought to the laboratory, and kept in
a container filled with running sea-water.
On 10 March 1996, about 250 larvae
hatched from one of the females. Of these,
60 were individually reared in 50 ml glass
bottles placed in an incubator at 9.5—
10.5°C. Some larvae were kept in 11 glass
beakers for mass culture.
Larvae were fed with newly hatched Ar-
temia nauplii. Moulting and mortality were
checked daily. After checking, the larvae
were transferred to freshly prepared bottles
and beakers. At each developmental stage,
dead larvae and exuviae were fixed and pre-
served with 3% neutralized formalin solu-
tion. At least five specimens of each stage
were dissected in ethylene glycol for mi-
croscopic observation. Drawings were
made with the aid of a drawing tube.
VOLUME 113, NUMBER 1
Number of Larvae
Zoea 1
Zoea 2
Zoea 3
Zoea 4
Megalopa
Days after Hatching
Fig.l.
ditions of 9.5—10.5°C and 31.80—32.65%o.
Measurements taken were: carapace
length (CL), from the anterior tip of the ros-
trum to the postero-median margin of the
carapace; total length (TL), from the ante-
rior tip of the rostrum to the postero-median
margin of the telson excluding telson pro-
cesses. At least five specimens were mea-
sured using a calibrated ocular micrometer.
Results
Development and duration of the lar-
vae.—Cryptolithodes expansus passed
Table 1.—Cryptolithodes expansus Miers. Duration
of each larval stage reared at 9.5~10.5°C and
31.80~32.65%o.
Duration of larval stages (day)
Larval stages Mean Range n
Zoea I 4.8 3-7 47
Zoea II 8.9 6-18 42
Zoea III 8.2 6-17 29
Zoea IV 14.8 13-16 18
Megalopa 10.5 7-15 ,
Cryptolithodes expansus Miers. Survival and duration of larval stages reared under laboratory con-
through four zoeal stages and a megalopa
(Table 1, Fig. 1). Of the 60 larvae reared
individually, only three molted to the me-
galopa stage within approximately 37 days.
Descriptions
First Zoea.
Size: CL = 1.6—1.8 mm (mean 1.6 mm);
TL = 4.0—4.4 mm (mean 4.1 mm).
Duration: 3-7 days.
Color: dark brown color uniformly dif-
fused over the whole carapace.
Carapace (Fig. 2A, B): rostrum well-de-
veloped, tapering from broad base to acute
tip; ventral and posterior carapace margins
with submarginal furrow and raised rim;
postero-lateral margin smoothly rounded.
Eyes (Fig. 2A, B): sessile.
Abdomen (Fig. 2 A, B): narrow and slen-
der; 5 somites plus telson; 3rd—Sth somites
with pair of lateral spines; pair of minute
spines on postero-dorsal margin of 2nd—Sth
somites.
56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Telson (Fig. 2L): armed with 7 pairs of
marginal processes; outermost a naked
spine; 2nd a fine hair; 3rd—7th denticulate
spines.
Antennule (Fig. 2C): biramous; an en-
dopodal bud naked, a delineated exopod
with 10 aesthetascs.
Antenna (Fig. 2D): biramous; scaphocer-
ite of the protopod with 6 long plumose se-
tae and 2 minute naked setae; endopod na-
ked, tapering to sharp terminal point, a
spine adjacent to the endopod.
Mandible (Fig. 2E): stout and well armed
with median teeth; mandibular palp present
as a small bud.
Maxillule (Fig. 2F): coxal endite with 7
serrated and 1 naked spinnule; basial endite
with 7 stout denticulate spines and 2 minute
setae; endopod 3-segmented with 2+1+3
setae.
Maxilla (Fig. 2G): proximal and distal
lobes of coxal endite with 7 and 4 setae
respectively; proximal and distal lobes of
basial endite with 4 setae each; endopod un-
segmented with 3 subterminal and 5 ter-
minal setae; scaphognathite with 10 plu-
mose setae.
First maxilliped (Fig. 2H): coxa naked;
basis with 2, 2, 3, 3 setae, endopod 5-seg-
mented with 3, 2, 1, 2, 4+1 setae; exopod-
ite with 4 terminal plumose natatory setae.
Second maxilliped (Fig. 21): coxa naked;
basis with 1, 2 setae; endopod 4-segmented
with 2, 2, 2, 4+1 setae; exopod with 4 plu-
mose natatory setae.
Third maxilliped (Fig. 2J): biramous; en-
dopod unsegmented; exopod two-segment-
ed.
Pereiopods (Fig. 2K): elongate but not
functional appendages; first pair subchelate.
Second Zoea
Size: CL = 2.1—2.2 mm (mean 2.2 mm);
TL = 4.4-5.0 mm (mean 4.8 mm).
Duration: 6—18 days.
Carapace (Fig. 3A, B): slight increase in
size, but no change in armature.
Eyes (Fig. 3A, B): stalked and movable.
Telson (Fig. 3L): unchanged.
Antennule (Fig. 3C): similar to first zoea
exopod with 9 aesthetascs of varying size.
Antenna (Fig. 3D): exopod with 7 plu-
mose setae and | minute naked seta.
Mandible (Fig. 3E): some teeth added.
Maxillule (Fig. 3F): setation of endites
and endopod unchanged.
Maxilla (Fig. 3G): proximal and distal
lobes of coxal endite with 9 and 4 setae
respectively; proximal and distal lobes of
basial endite with 5 and 4 setae respective-
ly; endopod unsegmented, with 3 subdter-
minal and 5 terminal setae; scaphognathite
with 22 plumose setae.
First maxilliped (Fig. 3H): setation of ba-
sis unchanged; | plumose lateral seta added
on lIst—3rd segments of endopod; exopod
with 8 plumose natatory setae.
Second maxilliped (Fig. 31): setation of
basis unchanged; 1 plumose lateral seta
added to Ist to 3rd segments of endopod;
exopod with 8 plumose natatory setae.
Third maxilliped (Fig. 3J): biramous; en-
dopod bud with 3 setae; exopod with 8 plu-
mose natatory setae.
Pereiopods (Fig. 3K): unchanged in ar-
mature; slight increase in size.
Pleopods (Fig. 3B): rudimentary unira-
mous buds on somites 2nd—Sth.
Third Zoea
Size: CL = 2.2—3.2 mm (mean 2.6 mm);
TL = 4.9-5.9 mm (mean 5.4 mm).
Duration: 6—17 days.
Carapace (Fig. 4A, B): similar to second
zoea.
Telson (Fig. 4L): unchanged.
Antennule (Fig. 4C): exopod with 9
aesthetascs.
Antenna (Fig. 4D): endopod much longer
than scaphocerite.
Mandible (Fig. 4E): some teeth added.
Maxillule (Fig. 4F): setation and shape
unchanged.
Maxilla (Fig. 4G): proximal and distal
lobes of coxal endite with 8 and 4 setae
respectively; basial endite unchanged; en-
VOLUME 113, NUMBER 1 57
Fig. 2. Cryptolithodes expansus Miers. First zoea. A, Dorsal view; B, Lateral view; C, Antennule; D, An-
tenna; E, Mandible; K Maxillule; G, Maxilla; H, First maxilliped; I, Second maxilliped; J, Third maxilliped; K,
Pereiopods; L, Telson. Scale bars = 0.2 mm.
58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Cryptolithodes expansus Miers. Second zoea. A, Dorsal view; B, Lateral view; C, Antennule; D,
Antenna; E, Mandible; K Maxillule; G, Maxilla; H, First maxilliped; I, Second maxilliped; J, Third maxilliped;
K; Pereiopods; L, Telson. Scale bars = 0.2 mm.
VOLUME 113, NUMBER 1
dopod unsegmented, with 3 subterminal
and 4 terminal setae; scaphognathite with
24 plumose setae.
First maxilliped (Fig. 4H): setation un-
changed.
Second maxilliped (Fig. 41): setation un-
changed.
Third maxilliped (Fig. 4J): setation un-
changed.
Pereiopods (Fig. 4K): unchanged.
Pleopods (Fig. 4B): well developed but
not functional appendages.
Fourth Zoea
Size: CL = 3.0—3.4 mm (mean 3.2 mm);
TL = 6.1—6.9 mm (mean 6.4 mm).
Duration: 13—16 days.
Carapace (Fig. 5A, B): similar to third
zoea.
Telson (Fig. 5T): unchanged.
Antennule (Fig. 5C): similar to previous
stage.
Antenna (Fig. 5D): endopod 2-segment-
ed and much longer than exopod.
Mandible (Fig. 5E): some teeth added.
Maxillule (Fig. 5F): setation of coxal en-
dite and endopod unchanged; basial endite
with 8 stout denticulate spines and 2 setu-
les.
Maxilla (Fig. 5G): setation of coxal and
basal endite unchanged; scaphognathite
with 26 plumose setae.
First maxilliped (Fig. 5H): setation un-
changed.
Second maxilliped (Fig. 51): setation un-
changed.
Third maxilliped (Fig. 5J): more devel-
oped than in third zoea.
Pereiopods (Fig. 5K—O): uniramous; Ist
pair unchange; 2nd—4th with apical spine
except on Sth.
Pleopods (Fig. 5P—S): unchanged.
Megalopa
Size: CL = 2.4—2.5 mm (mean 2.5 mm);
TL = 3.6—3.8 mm (mean 3.7 mm).
Duration: 7—15 days.
Color: intense scarlet.
39
Carapace (Fig. 6A, B): triangular in dor-
sal view, with conical rostrum, convex in
transverse section; cervical groove distinct;
angle between lateral carapace and pleuron
acute, with well-marked keel projecting lat-
erally.
Abdomen (Fig. 6A, B, O): 6 somites plus
telson; 2nd—Sth segments with pleopods;
6th segment with 3 minute hairs on the dis-
tal margin.
Telson (Fig. 6O): naked; more elongate
and conical; twice as long as 6th abdominal
segment.
Antennule (Fig. 6C): biramous; lower ra-
mus 2-segmented with 0, 3+3 setae; upper
ramus 4-segmented with 0, 4, 3, 3 aesthe-
tascs plus 3 terminal setae.
Antenna (Fig. 6D): scale naked; flagel-
lum 8-segmented with 0, 2, 0, 0, 4, 1, 4, 4
setae.
Mandible (Fig. 6E): strongly chitinized,
smooth and not toothed as in zoeal stages,
with broad blade-like process; palp 2-seg-
mented.
Maxillule (Fig. 6F): endites with minute
spines; endopod unsegmented.
Maxilla (Fig. 6G): setae of coxal and ba-
sial endite reduced and tooth-like; endopod
unsegmented; enlarged scaphognathite with
47 plumose setae.
First maxilliped (Fig. 6H): basis with 8
setae; endopod unsegmented and with 5 se-
tae; exopod with 4 terminal setae.
Second maxilliped (Fig. 61): basis with 2
setae; endopod 4-segmented with 0, 0, 0, 2
setae; exopod 2-segmented with 0, 4 setae.
Third maxilliped (Fig. 6J): basis with 1, 1
setae; endopod 5-segmented with 4, 6, 5, 14,
9 setae; exopod with 3 long plumose setae.
Pereiopods (Fig. 7A—E): well developed
and armed with spines; functional cheli-
peds.
Pleopods (Fig. 6K—N): present on 2nd—
5th abdominal somites; endopod small and
naked; exopods with 9-11 plumose nata-
tory setae.
Discussion
Morphological characteristics of lithodid
larvae have been discussed by Gurney
60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Cryptolithodes expansus Miers. Third zoea. A, Dorsal view; B, Lateral view; C, Antennule; D,
Antenna; E, Mandible; K Maxillule; G, Maxilla; H, First maxilliped; I, Second maxilliped; J, Third maxilliped;
K, Pereiopods; L, Telson. Scale bars = 0.2 mm.
(1942), MacDonald et al. (1957), Pike & to that of the Paguridae, except for the re-
Williamson (1960), Kurata (1964), and duction or disappearance of uropods.
Konishi (1986). They implied that larval Although the genus Cryptolithodes be-
morphology of this family was very similar longs to the family Lithodidae, it differs
VOLUME 113, NUMBER 1 61
Fig. 5. Cryptolithodes expansus Miers. Fourth zoea. A, Dorsal view; B, Lateral view; C, Antennule; D,
Antenna, D,-D,, Endopods of antenna; E, Mandible; F Maxillule; G, Maxilla; H, First maxilliped; I, Second
maxilliped; J, Third maxilliped; K, First pereiopod; L, Second pereiopod; M, Third pereiopod; N, Fourth pe-
reiopod; O, Fifth pereiopod; P, Pleopod of second abdominal segment; Q, Pleopod of third abdominal segment;
R, Pleopod of fourth abdominal segment; S, Pleopod of fifth segment; T, Telson. Scale bars = 0.2 mm.
62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 6. Cryptolithodes expansus Miers. Megalopa. A, Dorsal view; B, Lateral view; C, Antennule; D, An-
tenna; E, Mandible; K Maxillule; G, Maxilla; H, First maxilliped; I, Second maxilliped; J, Third maxilliped; K,
Pleopod of second abdominal segment; L, Pleopod of third abdominal segment; M, Pleopod of fourth abdominal
segment; N, Pleopod of fifth abdominal segment; O, Telson. Scale bars = 0.2 mm.
VOLUME 113, NUMBER 1 63
“i oe ; ea =
ee
vee Se
Fig. 7. Cryptolithodes expansus Miers. Megalopa. A, First pereiopod; B, Second pereiopod; C, Third pe-
reiopod; D, Fourth pereiopod; E, Fifth pereiopod. Scale bars = 0.2 mm.
Table 2.—Comparison of morphological characters between C. typicus and C. expansus
Species
C. typicus C. expansus
Characters (Hart 1965) (Present study)
First Zoea
Size* carapace length 1.4 mm 1.6 mm
total length 3.0 mm 4.1 mm
Antennule aesthetascs 8 10
Antenna endopod sharp tip, minute subterminal tooth. _ sharp tip
Maxillule basial endite 7 setae 9 setae
endopod TVisetae 9 setae
Maxilla proximal lobes 8, 6 setae 7, 4 setae
Telson posterior margin round straight
Second Zoea
Antennule aesthetascs 8 2)
Antenna endopod segmented unsegmented
Maxillule basial endue 8 setae 9 setae
endopod 1-1-3 setae 24715-3' setae
Third Zoea
Antenna endopod segmented unsegmented
Fourth Zoea
Antenna endopod segmented segmented
Maxillule basial endue 9 setae 10 setae
Megalopa
Size* Carapace length 2.0 mm 2.5 mm
Total length 2.9 mm 3.7 mm
Carapace rostrum wide and flattened narrow and pointed
Abdomen 5th segment 2 spines 3 spines
Antenna flagellum 2+1+ 1+-04+-3+-44+2+7 setae 0+2+0+0+4+1+4+4 setae
Third maxil-
liped exopod 4 plumose setae 3 plumose setae
Telson shape triangular spaculate
* Measured from Hart (1965, fig. 1 I, fig. 3)
64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
morphologically from other lithodid larvae.
The larvae of Cryptolithodes spp. resemble
those of some Diogenidae and Coenobitidae
in that they lack lateral carinae on the pos-
tero-lateral margins of the carapace. Major
morphological characteristics of lithodid
larvae as reported by Konishi (1986) are as
follows: ‘‘Uropods, if present, lack an en-
dopod, even in the final zoeal stage; the
third maxilliped has an endopod in the first
zoeal stage; abdominal somites lack medio-
dorsal spines; the telson without an anal
spine.”
Comparative morphological features of
Cryptolithodes typicus and C. expansus are
summarized in the Table 2. The larvae of
C. expansus differed from those of C. typ-
icus in body size, number of antennular
aesthetascs, setation of the maxillule, and
segmentation of the second maxilliped in
the zoeal stages.
The zoeal stages of Cryptolithodes typi-
cus and C. expansus differ from those of
the other lithodid larvae in the following:
The abdomen is narrower and more slender,
and composed of five segments plus telson
in all the zoeal stages; uropods are absent
in all the zoeal stages; a postero-lateral ca-
rinae on the carapace is absent in zoeal
stages; a mandibular palp is present at the
first zoeal stage.
Acknowledgements
We are very grateful for the helpful com-
ments of Drs. D. I. Williamson (Port Erin
Marine laboratory, Isle of Man, England)
and C. Sankarankutty (Universidade Fed-
eral do Rio Grande do Norte, Natal, Brazil).
We thank Mrs. D. N. Kim for the collection
of the ovigerous females. We also thank
Patsy A. McLauglin (Shannon Point Marine
Center, Washington, U.S.A.) and two other
anomymous reviewers for their valuable
criticisms and suggestions in improving the
manuscript.
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Sars, G. O. 1890. Bidrag Kundskaben om Decapod-
ernes Forvandlinger. II: Lithodes-Eupagurus-
Spiropagurus-Galathodes-Munida-Porcellana-
(Nephros).—Archiv for Mathematik og Naturv-
idenskab 13:133-—201.
Schalfeew, P. 1892. Carcinologische Bemerkungen aus
dem Zoologischen Museum der Kaiserlichen
Akademie der Wissenschaften.—Bulletin de
lV’ Académie Impériale des Sciences de Saint-Pé-
tersbourg 35:33 1-342.
Tilesius, W. C. 1815. De cancris camtschaticis, oniscis,
entomostracis et cancellis marinis microscopicis
noctilucentibus, cum tabulis IV: aenaeis et ap-
pendice adnexo de acaris et ricinis Camtscha-
ticis. Conventui exhibuit die 3 Februarii
1813.—Mémoires de |’ Académie Impériale de
Sciences de St Pétersbourg, 5:331—405.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):66—69. 2000.
A new genus of pinnotherid crab from the Indian Ocean
(Crustacea: Decapoda: Brachyura)
Raymond B. Manning and Bella Galil
(RBM) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560-0163, U.S.A.; (BG) Israel Oceanographic and Limnological
Research Ltd., RO. Box 8030, Haifa 31080, Israel
Abstract.—Abyssotheres, new genus, is recognized for Pinnotheres abyssi-
cola Alcock & Anderson, 1899, known from a single female taken in a bivalve
shell from a depth of 787 m off Travancore, India. In this new genus the dactyli
of walking legs 1 and 2 are longer than those of walking legs 3 and 4. This is
the eighth genus of pinnotherids that shares a two-segmented palp on the third
maxilliped.
Alcock & Anderson (1899) described
Pinnotheres abyssicola from a single ovig-
erous female found in a lamellibranch taken
off Travancore, India at a depth of 430 fm
(787 m). It is known only from the holotype
kept in the Zoological Survey of India and
remains the deepest recorded occurrence of
a pinnotherid crab.
One of us (BG) recently visited the Zoo-
logical Survey of India (formerly the Indian
Museum) and examined the holotype of P.
abyssicola. It is in very poor condition, pos-
sibly having dried out. However, she was
able to examine the third maxilliped (Fig.
la) and determined that the palp comprises
only two segments. Campos (1996) re-
viewed the pinnotherid genera with a two-
segmented palp on the third maxilliped and
reported that the two-segmented palp was
found only on members of six genera. Man-
ning (1993) added a seventh, Epulotheres.
A combination of characters distinguishes
P. abyssicola from all pinnotherid genera
with a two-segmented palp. We recognize a
new genus here for it.
We use the following abbreviations in the
account below: fm, fathom(s); m, meter(s);
MXP3, third maxilliped; WL, walking leg(s).
Abyssotheres, new genus
Diagnosis.—Size medium, carapace
length and width described as less than 10
mm in adult. Carapace length and width
subequal, front prominent, transverse, pro-
jecting anteriorly beyond eyes. Eyes visible
in dorsal view. MXP3 with ischium and
merus indistinguishably fused, arched, in-
ner margin projecting at about distal third.
Palp 2-segmented (Fig. la), terminal seg-
ment spatulate, shorter than preceding seg-
ment. Chela with dactylus slightly less than
half of propodus. Walking legs (Fig. 1b)
slender, equal right and left; WL1-2 with
dactyli longer than dactyli of WL3-4. Ab-
domen unknown.
Male.—Unknown.
Type species.—Pinnotheres abyssicola
Alcock & Anderson, 1899, by present des-
ignation and monotypy.
Etymology.—From the Latin abyss,
depth, and the ending theres.
Host.—A large bivalve, Acesta indica
(Smith) (originally described in Lima) (AlI-
cock & Anderson 1899).
Distribution.—Known only from off the
coast of Travancore, India, at a depth of 787
m. This is the deepest record for a pinnoth-
erid.
Remarks.—Alcock & Anderson’s origi-
nal account, based on an ovigerous female
8 mm wide, is: “‘Carapace as long as broad,
circular, smooth; front rather prominent,
about one-fifth the greatest breadth of the
VOLUME 113, NUMBER 1
67
es
Eig: I.
mim: a MXP3; b, WL.
carapace. The whole of the eyes and eye-
stalks and almost the whole of the orbit are
visible in dorsal view. The eyes are well
developed, but very pale. The dactylus of
the external maxillipeds is styliform and is
inserted at the end of the preceding joint.
The lower border of the thumb is fringed
with fine hairs. The legs are slender; the
second and third pair are both about 1%
times as long as the carapace, and have the
dactylus slightly longer than it is in the oth-
er two pairs” (Alcock & Anderson 1899:
14).
This account provides few diagnostic
features. First, the carapace is round, as
broad as long, and the front is fairly prom-
inent.
Second, the terminal segment of the
MXP3 palp is articulated terminally on the
subdistal segment. Among those Pinnoth-
erinae with a two-segmented palp, the dac-
tylus of the palp is articulated terminally in
members of Orthotheres Sakai, 1969 (see
Campos 1989, Manning 1993) and in three
species described by Birger (1895), Pin-
notheres glaber, P. impressus, and P. lae-
Abyssotheres abyssicola (Alcock & Anderson, 1899). Ovigerous female holotype, carapace length 8
vis, all presumably shore species known
from the Pacific Ocean. In Biirgers’s species
the dactyli of the walking legs are equally
long.
A third distinguishing character of A.
abyssicola is that the dactyli of WL1-2 are
longer than those of WL3-4. This may be
an unique feature within the Pinnotheridae.
A fourth characteristic feature of A. abys-
sicola is the arched MXP3, a feature shared
with the unrelated Limotheres nasutus Hol-
thuis, 1975, from the Caribbean. It also is
a commensal of a species of Lima, but oc-
curs in shallow water. Limotheres has a
three-segmented mandibular palp, and dif-
fers from Abyssotheres in numerous other
features.
Campos (1996) studied six genera of pin-
notherids that have a two-segmented palp
on the MXP3: Calyptraeotheres Campos,
1990; Dissodactylus Smith, 1870; Gem-
motheres Campos, 1996; Ostracotheres H.
Milne Edwards, 1853; Tunicotheres Cam-
pos, 1996; and Xanthasia White, 1846.
Members of Abyssotheres can be distin-
guished from members of these genera as
68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
follows: the carapace lacks the sharp lateral
margins and the dorsal sulci characteristic
of Calyptraeotheres; the apices of WL dac-
tyli are simple, rather than bifurcated as in
Dissodactylus; MXP3 with the inner distal
margin of the merus obtusely angled rather
than curved as in Gemmotheres, Ostracoth-
eres, and Tunicotheres; carapace lacking
upturned lateral margins and median mush-
room-shaped tubercle as in Xanthasia. All
of these other genera comprise shore rather
than slope species, and all but Ostracoth-
eres and Xanthasia are restricted to the
Americas.
A seventh genus, Epulotheres, was added
by Manning (1993), who incorrectly re-
ported that it had a MXP3 with a three-
segmented palp. This was corrected by
Manning & Felder (1996) who reillustrated
the palp, which has the usual three seg-
ments. Nannotheres Manning & Felder,
1996 does have a two-segmented palp. In it
WL4 is the longest walking leg and the dac-
tyli of the walking legs are similar and
equal in length.
The obtuse projection on the dorsal sur-
face of the dactylus of the WL (Fig. 1b)
may prove to be an unique feature of A.
abyssicola.
Only one other pinnotherid, Alain cros-
nieri Manning, 1998, is known from depths
greater than 400 m. An associate of holo-
thurians, it was taken in depths of 399-461
m off Indonesia.
We are pleased to have the opportunity
to include this report in a volume dedicated
to our late colleague Austin B. Williams,
whose many studies on decapods have con-
tributed much to our knowledge of the
group.
Acknowledgments
We acknowledge with thanks the coop-
eration of the Director, Zoological Survey
of India, Calcutta, for permission for one of
us (BG) to examine material in the care of
that institution. The figure was prepared by
Lilly King Manning. We thank Ernesto
Campos and Roy K. Kropp for their very
helpful reviews of the manuscript. Man-
ning’s studies of pinnotherid crabs are sup-
ported by the Smithsonian Marine Station
at Fort Pierce. This is contribution no. 487
from that facility.
Literature Cited
Alcock, A., & A. R. S. Anderson. 1899. An account
of the deep-sea Crustacea dredged during the
survey season of 1897-98. Natural History
Notes from H. M. Royal Indian Marine Survey
Ship “Investigator,” Commander T. M. Heming,
R. N., commanding, series 3, no. 2.—Annals
and Magazine of Natural History, series 7, 3:1—
27, 278-292.
Birger, O. 1895. Ein Beitrag zur Kenntniss der Pin-
notherinen.—Zoologische Jahrbiicher, Abthei-
lung fiir Systematik, Geographie und Biologie
der Thiere 8:361—390.
Campos, E. 1989. Comments on taxonomy of the ge-
nus Orthotheres Sakai 1969 (Crustacea, Deca-
poda, Pinnotheridae).—Bulletin of Marine Sci-
ence 44:1123-—1128.
. 1990. Calyptraeotheres, a new genus of Pin-
notheridae for the limpet crab Fabia granti
Glassell, 1933 (Crustacea, Brachyura).—Pro-
ceedings of the Biological Society of Washing-
ton 106:92-101.
. 1996. Partial revision of pinnotherid crab gen-
era with a two-segmented palp on the third
maxilliped (Decapoda: Brachyura).—Journal of
Crustacean Biology 16(3):556—563.
Holthuis, L. B. 1975. Limotheres, a new genus of pin-
notherid crab, commensal of the bivalve Lima,
from the Caribbean Sea.—Zoologische Mede-
delingen 48(25):291—295.
Manning, R. B. 1993. West African pinnotherid crabs,
subfamily Pinnotherinae (Crustacea, Decapoda,
Brachyura).—Bulletin du Muséum national
d’ Histoire naturelle, Paris, série 4, 15 (A, 1-4):
125-177.
. 1993. Epulotheres angelae, new genus, new
species, a pinnotherid crab from the Caribbean
Sea (Decapoda: Pinnotheridae).—Journal of
Crustacean Biology 13(4):801—804.
. 1998. A new genus and species of pinnotherid
crab (Crustacea, Decapoda, Brachyura) from In-
donesia.—Zoosystema 20(2):357-—362.
, & D. L. Felder. 1996. Nannotheres moorei, a
new genus and species of minute pinnotherid
crab from Belize, Caribbean Sea (Crustacea:
Decapoda: Pinnotheridae).—Proceedings of the
Biological Society of Washington 109(2):311-—
oh We
Milne Edwards, H. 1853. Mémoire sur la famille des
VOLUME 113, NUMBER | 69
Ocypodiens, suite.—Annales des Sciences Na- ican Crustacea, I.—Transactions of the Con-
turelle, série 3, Zoologie, 20:163—228. necticut Academy of Arts and Sciences 2:1] 13—
Sakai, T. 1969. Two new genera and twenty-two new 176.
species of crabs from Japan.—Proceedings of the | White, A. 1846. Notes on four genera of Crustacea.
Biological Society of Washington 82:243—280. Annals and Magazine of Natural History 18:
Smith, S. I. 1870. Ocypodidea. Notes on North Amer- 176-178.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):70-—76. 2000.
A new genus and species of ghost shrimp from Tobago, West Indies
(Crustacea: Decapoda: Callianassidae)
Richard Heard and Raymond B. Manning
(RH) Gulf Coast Research Laboratory, East Beach Road, Ocean Springs, Mississippi 39564,
U.S.A.; (RBM) Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A.
Abstract.—The new genus Pseudobiffarius is the seventh genus of Calli-
anassinae recognized from the Americas. This monotypic genus is based on its
type species from Tobago, West Indies, Pseudobiffarius caesari, new species.
It differs from all members of the subfamily in having a very short, stout upper
flagellum on the Al peduncle.
Collections of a very small callianassid
were made on Tobago by two individuals
at different times, Lois Nickell in 1989 and
Richard Heard in 1992 and 1993. The spe-
cies was initially identified as a member of
Biffarius Manning & Felder, 1991, but clos-
er examination after Heard’s collections of
1993 revealed that it represented not only a
new species but also a new genus as well.
The new taxa are named here.
The types have been deposited in the Na-
tional Museum of Natural History, Smithson-
ian Institution, Washington, D.C. (USNM).
Carapace length (cl) measured as post-
orbital carapace length and total length (t1),
measured on the midlineare in millimeters
(mm). Other abbreviations: Al, antennule;
A2, antenna; leg., collector; m, meter(s);
Max1-—2, maxillae 1—2; Mxp1-3, maxilli-
peds 1—3; P1—5, pereopods 1—5 (P1 and P2
are chelipeds, P3—5 walking legs); Plp1—S,
pleopods 1-5.
Family Callianassidae Dana, 1852
Subfamily Callianassinae Dana, 1852
Pseudobiffarius, new genus
Type species.—Pseudobiffarius caesari,
new species, by present designation and
monotypy.
Etymology.—The generic name _ is
formed by combining the Latin prefix pseu-
do-, false, with the generic name Biffarius,
alluding to the resemblance of the present
genus to the latter. The gender is masculine.
Diagnosis.—Size relatively small, tl of
adults less than 30 mm. Carapace with acute
rostral projection, lacking rostral carina or
spine. Cornea dorsal, subterminal, disk-
shaped, corneal elements distinct. Al pedun-
cle longer than A2 peduncle; dorsal flagel-
lum of Al peduncle short and stout, shorter
than peduncle. Mxp3 without exopod, ischi-
um-merus operculiform; inner face of ischi-
um with cristate ridge of strong teeth; merus
projecting slightly beyond articulation with
carpus; last 3 segments slender. Chelipeds
unequal in male, equal in female, with meral
hook. Plpl uniramous in both sexes; Plp2
absent in male, biramous in female; Plp3—5
foliaceus, with stubby, projecting appendices
internae in both sexes.
Remarks.—Manning & Felder (1991)
recognized and named three American gen-
era in this subfamily, Biffarius, Neotrypaea,
and Notiax, and in 1992 added a fourth,
Gilvossius. Rodrigues & Manning (1992)
added a fifth, Poti. Heard & Manning
(1998) added a sixth, Necallianassa. Poti
and the type genus of the family, Calli-
anassa Leach, 1814, differ from the re-
mainder of these genera by having slender
Mxp3, which are pediform rather than op-
erculiform; Poti also differs from all other
VOLUME 113, NUMBER 1
genera in the subfamily by having an in-
complete linea thalassinica on the carapace.
Members of Neotrypaea and Notiax are
known only from the eastern Pacific; mem-
bers of the other genera are known only
from the western Atlantic.
Members of Pseudobiffarius can be dis-
tinguished at once from the American gen-
era with broad Mxp3 by the short, stout dor-
sal flagellum of Al. They further differ from
both Neotrypaea and Gilvossius in having
the appendix internae of the Plp3—5 embed-
ded in the edge of the pleopod, whereas they
are projecting in Pseudobiffarius, as in Bif-
farius, Necallianassa and Notiax. In Notiax
the second male pleopod is present, whereas
it is absent in Pseudobiffarius; members of
Notiax also have a strong rostral spine ex-
tending almost beyond the cornea and a me-
dian distal spine on the telson. Members of
Biffarius lack the strong ridge of teeth pre-
sent on the inner margin of the Mxp3 in
members of Pseudobiffarius.
Pseudobiffarius caesari, new species
Figs. 1, 2, 3a—k, n—o, 4, 5a
Material.—Tobago: Lover’s Beach,
northwest corner of Man O’War Bay
[11°19’N, 60°34’W], protected beach and
Shallow reef, depth ca. 2 m, leg. R. Heard,
sta 3, 6 Apr 1992: 1 male, cl 3.2 mm (par-
atype, USNM 260965).
Pirate’s Cove, east side of Man O’ War
Bay, depth 2—3 m, leg. R. Heard, 12 Jan
1992: 3 females, 2 non-ovigerous, cl 2.6
and 3.8, 1 ovigerous, cl 4.1 (paratypes,
USNM 260970).
Buccoo Reef [11°11’N, 60°49’W], back
reef area with sand bottom, depth ca. 2 m,
leg. R. Heard, sta 10A (1992 sta 2), 11 Jan
1993: 1 male, cl 4.7 mm (holotype, USNM
260966).
Coral Gardens, Buccoo Reef, depth 5 m,
leg. Lois Nickell, 6 Jul 1989: 1 female, cl
3.6 mm (paratype, USNM 260968).—Coral
Gardens, Buccoo Reef, depth 2 m, leg. Lois
Nickell, 2 Aug 1989: 1 female, cl 6.0 mm,
tl 26 mm (paratype, USNM 260969).
71
Pigeon Point [11°10’N, 60°51’W], depth
1 m, on sand, leg. R. Heard, 15 Jan 1993:
1 female, cl 4.4 mm (paratype, USNM
260967).
Lowlands Lagoon [= Petit Trou;
13°50'N, 61°05’W], leg. R. Heard, sta. 9, 7
Apr 1992: 1 male, cl 5.4 mm (paratype,
USNM 260971).
Diagnosis.—Size very small, cl 6.0 mm
or less and tl 30 mm or less in adults. Tel-
son subrectangular, with small submarginal
spines posteriorly and distinct median and
posterolateral spines. Mxp3 ischium-merus
operculiform, without exopod, inner surface
of ischium with strong crest of teeth. Major
cheliped not dimorphic in males; merus of
both chelipeds with ventral hook. Male
lacking Plp2. Uropods unarmed.
Description.—Carapace (Figs. 1, 4a—c)
smooth, with dorsal oval, without cardiac
prominence; cervical groove distinct; linea
thalassinica distinct, parallel to longitudinal
axis of body; rostrum short, acute, not pro-
duced into distinct spine; lateral frontal pro-
jections present, low, obtuse.
Abdominal somites (Figs. 1, 4a, d)
smooth, somite 1 saddle-like, shortest; so-
mite 2 longest, almost twice as long as so-
mite 6, with small tuft of setae posterolat-
erally, set in submarginal longitudinal
crease; somites 3-5 subequal in length,
each with small lateral row of setae set in
submarginal crease; somite 6 about as long
as telson, with median carina. Telson (Fig.
4e, f) subrectangular, tapering posteriorly,
slightly longer than wide, with anterior tuft
of setae mid-dorsally and some isolated se-
tae and small spinules dorsally, submarginal
row of spinules present posteriorly; poste-
rior margin setose, emarginate, with median
spine and 2 pairs of posterolateral spines.
Eyes (Figs. 1, 4a—c, 5a) flattened, slightly
convex, about as long as first segment of
Al peduncle, external margins moderately
convergent, mesial margins parallel, tips
pointed; cornea rounded, darkly pigmented,
subterminal.
Al peduncle (Figs. 4a, b, 5a) extending
beyond A2 peduncle; segment 3 more than
72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figi
Reef. Lateral view. Scale = 1 mm.
twice as long as segment 2, segment 1 sub-
equal to segment 2; upper flagellum stout,
much shorter than distal segment of pedun-
cle. A2 peduncle (Figs. 4a, b, 5a) with seg-
ment 4 slightly shorter than segment 3 and
as long as segment | and 2 together; anten-
nal scale not evident; flagellum much lon-
ger than peduncle.
Mandible (Fig. 3a, b) with numerous
small teeth on incisor process; molar pro-
cess unarmed; palp 3-segmented, segment
3 elongate, longer than segments 1 and 2
combined, tapering distally. Max1 (Fig. 3c)
with broad basal endite, distal lobe almost
forming right angle; coxal endite elongate,
triangular; palp slender, with curved tip.
Max2 (Fig. 3d) with protopodal endites bi-
lobed, distalmost lobe elongate, triangular;
palp slender; exopod broad, with crenulated
margins. Mxp1 (Fig. 3e) with protopodal
endites very slender, bilobed; palp very
small but distinct, longer than wide; exopod
bilobed, mesial margin densely setose with
short, spatulate setae (Fig. 3f); epipod sub-
rectangular, elongate. Mxp2 (Fig. 3g) with
Pseudobiffarius caesari, new genus and species, male holotype (USNM 260966), cl 4.7 mm, Buccoo
5-segmented endopod, terminal segment
short, spatulate; exopod unsegmented,
shorter than endopodal merus; protopod
with small setose epipod, apparently lack-
ing arthrobranch. Mxp3 (Fig. 3h—k) ischi-
um-merus sub-operculiform; ischium about
as wide as long, mesial surface with dentate
crest of large teeth; merus more than 1.5
times wider than long, almost 4 times wider
than carpus and propodus, distal margin of
outer face projecting beyond articulation
with carpus; propodus much longer than
wide, tapering distally, only slightly wider
than digitiform dactylus.
Males with very unequal chelipeds. In
larger cheliped of male (Figs. 1, 2a, b): is-
chium with ventral margin spinulose in
largest of 3 males; merus with dorsal mar-
gin spinulose in proximal fourth, smooth
distally, posterior margin with spinules
proximal to distally-directed hook, with dis-
tal portion serrated; carpus distinctly longer
than wide, smooth dorsally and ventrally,
broadly rounded proximally; palm shorter
than carpus, length less than height; fingers
VOLUME 113, NUMBER 1
73
Fig. 2. Pseudobiffarius caesari, new genus and species. a, Male major Pl, outer aspect; b, Male major P1,
inner aspect; c, Female P2; d, Male P2; e, Male minor P1; f, Juvenile male minor P1, setae omitted; g, Juvenile
male major Pl; h, Male P3; i, Female P4; j, Female P5; k, Female Plp1; 1, Female Plp2. a, b, d, male holotype
(USNM 260966), cl 4.7 mm, Buccoo Reef; c, g, h, k, 1, female paratype (USNM 260967), cl 4.4 mm, Pigeon
Point; e, f, juvenile male paratype (USNM 260965), cl 3.2 mm, Lover’s Beach. Scale: a—j, 0.5 mm; k, 1, 1.0 mm.
about as long as palm, gaping, tips acutely
pointed, crossing; cutting edge of movable
finger minutely serrated; gape conspicuous,
base with dorsal convex lobe, edge serrate,
above smoother, deep invagination in distal
border of propodus.
Smaller cheliped of male (Fig. 2d, e) and
both chelipeds of female (Fig. 2c) similar in
size (chelipeds symmetrical in female): is-
chium smooth, unarmed, about as long as
merus; merus with small, acute hook at mid-
length of ventral margin; carpus longer than
Table 1.—Gill formula of Pseudobiffarius caesari,
new genus and species.
Maxillipeds Pereopods
vie. 3 TP a eee ee
Pleurobranchs = 2 aap AeA LE ES.
Arthrobranchs - — 2 aS 2 Qe
Podobranchs ey pees ee. eee
Epipods a aa
Exopods ee Se
other joints, length about 2.5 times height;
palm less than half as long as merus, length
and height subequal; fingers subequal,
curved, slightly longer than palm, sharply
pointed, with some serrations on cutting
edge of fixed finger, tip of fingers crossing.
P2 (Fig. 2c, d) with fingers not gaping, cut-
ting edges straight, smooth. P3 (Fig. 2h) with
ischium and merus of about same width, mer-
us longer than ischium; carpus broadening
distally; propodus wider than carpus, anterior
and posterior margins convex, posterior mar-
gin much longer than anterior; dactylus as
long as wide, spatulate. P4 (Fig. 2j) lacking
subchelar projection. PS (Fig. 21) distinctly
chelate, fingers small, tips curved.
Branchial formula, differing from normal
callianassid formula in complete absence of
arthrobranch on Mxp2, as shown in Table 1.
Plpl of male uniramous, 2-segmented.
Male lacking Plp2. Plp1 of female (Fig. 2k)
uniramous, 2-segmented, proximal segment
74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig: 3:
Pseudobiffarius caesari, new genus and species: a—k, n, 0, male holotype (USNM 260966), cl 4.7
mm, Buccoo Reef. Pseudobiffarius caesari, new genus and species: h, i, female paratype (USNM 260968), cl
3.6 mm, Buccoo Reef.—Biffarius biformis (Biffar, 1971): 1, m, male (USNM 266071), cl 5.5 mm, Fort Pierce,
Florida (USNM 266071): a—b, Mandible; c, Max1; d, Max2, e, Mxp1; f, Marginal setae of Mxpl1, enlarged; g,
Mxp2; h, Mxp3, inner face; i, Mxp3, outer face; j, Mxp3, inner face); 1, Mxp3, outer face; m, Mxp3, inner face;
n, 0, Plp3 appendix interna. Upper scale, a—e, g-m = 0.5 mm; lower scale, f, n, o, = 1.0 mm.
with subterminal and terminal patch of short
setae; distal joint about as long as proximal,
with a patch of small setae at midlength and
4 spiniform apical setae. Plp2 of female
(Fig. 21) biramous; endopod straight, with
long terminal setae and a few setae scattered
along its length; exopod similar to endopod
but curved and lacking longer apical setae.
Plp3—5 with stubby, projecting appendices
internae (Fig. 3n, 0) in both sexes.
Uropodal endopod (Fig. 4a, e) slightly
longer than wide, margins densely setose;
exopod slightly longer than and much wider
than endopod, upper plate more than half as
wide as lower.
Size.—cl of males (n = 3), 3.2 to 5.4 mm;
tl of largest male, 20 mm; cl of females (n
= 6), 2.6 to 6.0 mm, of ovigerous female,
4.1 mm; tl of largest female, 26 mm.
Remarks.—Although this species superfi-
VOLUME 113, NUMBER 1
75
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Fig. 4. Pseudobiffarius caesari, n. gen., n. sp. a, Dorsal view (pereopods omitted); b, Carapace and anterior
appendages, dorsal view; c, Carapace and anterior appendages, lateral view; d, Abdomen, lateral view; e, Sixth
abdominal somite, telson, and left uropod, dorsal view; f, Telson, dorsal view, enlarged; g, Details of telson
margin, enlarged. a—d, f—g, female paratype (USNM 260967), cl 4.4 mm, Pigeon Point; e, female paratype
(USNM 260968), cl 3.6 mm, Buccoo Reef. Scale: a, c-e, 2 mm; b, f, 1 mm; g, 0.5 mm.
cially resembles the type species of Biffarius,
B. biformis (Biffar 1971), it differs in many
features. In contrast to B. biformis, males
have only one type of cheliped rather than
two types, as reported by Biffar (1971) in B.
biformis. Pseudobiffarius caesari also differs
from B. biformis in having the A2 peduncle
shorter than the Al peduncle, the upper Al
flagellum much shorter, stouter, and with 9 or
10 rather than 18 to 20 segments. In B. bi-
formis the inner face of the ischium of Mxp3
(Fig. 3m) lacks spinules, whereas there is a
line of strong spinules on the inner surface of
Mxp3 (Fig. 3h, j) in P. caesari.
Examination of the telson under high
magnification (Fig. 4f, g) yielded a very
different picture of its marginal ornamen-
tation than can be obtained under the mag-
nification available from a dissecting micro-
scope. In addition to being lined with short,
plumose setae and some much longer ones,
especially laterally, the posterior margin of
the telson has a distinct median spine and
a series of small, thick marginal spines and
shorter submarginal spines, with two larger
spines posterolaterally.
The shore fauna of Tobago is very rich
in species of burrowing decapods. Among
76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Pip...
female paratype (USNM 260967), cl 4.4 mm, Pigeon Point; b, Biffarius biformis (Biffar 1971), male, (USNM
266071, cl 5.5 mm, Fort Pierce, Florida. Scale = 1 mm.
the new species discovered by us (RH) are
another callianassid, Neocallichirus nickel-
lae Manning, 1993 and the commensal pin-
notherid crab, Austinixa hardyi Heard &
Manning, 1997.
We are pleased to acknowledge our re-
spect for our late colleague, Austin B. Wil-
liams, by being able pubiish this report in
a volume of the Proceedings of the Biolog-
ical Society of Washington dedicated to
him.
Etymology.—Named for Errol Caesar of
the Fisheries Division, Ministry of Agri-
culture, Land, and Marine Resources, To-
bago. His enthusiastic support and interest
in Heard’s fieldwork on Tobago materially
improved collecting opportunities.
Acknowledgments
We thank both Errol Caesar and J. David
Hardy for their support of Heard’s investi-
gations on Tobago; Lois Nickell, then with
the University Marine Biological Station,
Miullport, Scotland, for the gift of her col-
lections; and Lilly King Manning for pre-
paring the figures. Manning’s studies on
callianassid systematics are supported by
the Smithsonian Marine Station at Fort
Pierce; this is contribution no. 455 from
that facility.
Carapace and anterior appendages in lateral view. a, Pseudobiffarius caesari, new genus and species,
Literature Cited
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capoda, Thalassinidea) from the western Atlan-
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Heard, R. W., & R. B. Manning. 1997. Austinixa, a
new genus of pinnotherid crab (Crustacea: De-
capoda: Brachyura), with the description of A.
hardyi, a new species from Tobago, West In-
dies.—Proceedings of the Biological Society of
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,& . 1998. A new genus and species of
ghost shrimp (Crustacea: Decapoda: Callianas-
sidae) from the Atlantic Ocean.—Proceedings
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Leach, W. E. 1814. Crustaceology. /n D. Brewster, ed.,
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Manning, R. B. 1993. Two new species of Neocalli-
chirus from the Caribbean Sea (Crustacea: De-
capoda: Callianassidae).—Proceedings of the
Biological Society of Washington 106:106—114.
, & D. L. Felder. 1991. Revision of the Amer-
ican Callianassidae (Crustacea: Decapoda:
Thalassinidea).—Proceedings of the Biological
Society of Washington 104:762—790.
sates . 1992. Gilvossius, a new genus of
callianassid shrimp from the eastern United
States (Crustacea: Decapoda: Thalassinidea).—
Bulletin of Marine Science 49(1—2)[for 1991]:
558-561.
Rodrigues, S. de A., & R. B. Manning. 1992. Poti
gaucho, a new genus and species of ghost
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ence 51:9—13.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):77-87. 2000.
Griceus buskeyi, a new genus and species of calanoid copepod
(Crustacea) from benthopelagic waters off Hawaii
Frank D. Ferrari and E. L. Markhaseva
(FDF) Department of Invertebrate Zoology; National Museum of Natural History; Smithsonian
Institution; Washington, D.C. 20560-0534, U.S.A.; (ELM) Academy of Sciences, Zoological
Institute; Universitetskaya nab. 1; 199034 St. Petersburg, Russia
Abstract.—The new genus and species of deep-sea, benthopelagic calanoid
copepod, Griceus buskeyi, differs from other calanoid copepods in the follow-
ing derived character states: dorsal interlocking extensions of abdominal so-
mites; antenna 2, mandible, and maxilla 1 originate posterior to the labrum and
paragnaths; coxal endite of the mandible outside of the labral-paragnathal en-
velope; praecoxal and coxal endites of maxilla 1 unarmed; praecoxal endites
of maxilla 2 unarmed; endopod of maxilliped with four articulating segments.
Two setae on middle endopodal segment of swimming legs 3 and 4 suggests
the new genus and species belongs to the superfamily Augaptiloidea. A man-
dibular endopod which is reduced in size to a small unarmed lobe and absence
of an endopod on maxilla | suggests an affinity with the arietellid genus Par-
augaptilus Wolfenden.
Samples from deep-sea, benthopelagic
habitats continue to reveal a diverse fauna
of calanoid copepods which often express
unusual morphologies. Here we describe an
unusual species of calanoid which we be-
lieve can be placed in the family Arietelli-
dae as it is presently understood (Ohtsuka
et al. 1994), but not in any of its known
genera.
Methods
This adult female calanoid copepod was
collected 6 July 1997 from seawater flow-
ing from a flexible plastic pipe maintained
by Natural Energy Laboratory of Hawaii
Authority near Kona, Island of Hawaii,
19°43'27.01"N, 156°04'35.46”"W. The intake
of the pipe (1 m diameter) is located at 675
m, about 30 m from the bottom, and draws
in about 132 cubic m/min. Because of its
flexibility, the position of the intake may
vary vertically + 10 m with the movement
of the tide. Water from one outflow pipe (10
cm in diameter) drains through a 53 micron
mesh net placed in a large, dark tank. The
sample was collected after 12 hrs; the ani-
mal reported here may have been dead prior
to sample fixation with 4% formaldehyde.
The specimen was later cleared in steps
through 50% lactic acid/50% water to
100% lactic acid, stained by adding a so-
lution of chlorazol black E dissolved in
70% ethanol/30% water, and examined with
bright-field and with differential interfer-
ence optics.
Cephalic appendages are abbreviated Al
= antenna 1; A2 = antenna 2; Mn = man-
dible; Mx1 = maxilla 1; Mx2 = maxilla 2.
Appendages on thoracic somites are Mxp =
maxilliped (thoracopod 1); Pl-5 = swim-
ming legs (thoracopods 2—6). The caudal
ramus is CR. Designations of appendage
segments are according to Ferrari (1995) as
follows: medial lobe of a segment = hi, lat-
eral lobe = le; rami are exopod = Re and
endopod = Ri; ramal segments of Mx2 are
exopodal; the Mxp has a basopod with a
78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
distal medial lobe, and with at most five
endopodal segments in calanoids (Ferrari &
Dahms 1998). Armament elements of ap-
pendages are termed setae regardless of
their position or degree of rigidity. Setules
are epicuticular extensions of a seta; denti-
cles are epicuticular extensions of an ap-
pendage segment; spinules are epicuticular
extensions of a somite.
Calanoid superfamily names follow An-
dronov (1974) with the exception of Clau-
socalanoidea for Pseudocalanoida (Bow-
man 1987, International Commission on
Zoological Nomenclature 1988). The
changes proposed by Andronov (1991) are
not followed here because there is no re-
quirement that a family group name be de-
rived from the oldest included genus name,
or by coordination here the oldest included
family name (International Code of Zoolog-
ical Nomenclature 1985, articles 62—64).
The correct names in question are Augap-
tiloidea, Centropagoidea and Megacalano-
idea.
Griceus, new genus
Diagnosis.—The following derived char-
acter states are assumed to be shared by all
species of this genus discovered subse-
quently: dorsal interlocking extensions of
abdominal somites; A2, Mn and Mx! orig-
inating posterior to the labrum and parag-
naths; coxal endite of Mn outside of the la-
bral-paragnathal envelope; praecoxal and
coxal endites of Mx1 unarmed; praecoxal
endites of Mx2 unarmed; endopod of Mxp
with four articulating segments.
Type species.—Griceus buskeyi, by mon-
otypy.
Etymology.—The name honors Dr.
George D. Grice for his contributions to the
exploration of the deep-sea benthopelagic
fauna, and to the taxonomy of calanoid co-
pepods. Dr. Grice’s family name is latinized
to form Griceus. The gender of the name is
masculine.
Griceus buskeyi, new species
Specimens.—Holotype (National Muse-
um of Natural History, Smithsonian Insti-
tution USNM 288058) a dissected female,
0.90 mm in length; prosome—0.74 mm and
urosome—O.16 mm.
Female.—Pr (Fig. 1C): 6 segments; Ist a
complex of 5 cephalic somites plus Thl;
Th2—4 simple and articulated; arthrodial
membrane between Th5 and 6 incomplete;
reduced in width dorsally. Rostral area a
simple, bilobe plate (Fig. 1A, C) without
spinules or attenuations of cephalon. Lo-
cation of base of A2, Mn, Mx1, Mx2 and
Mxp as in Fig. 1A, B. Base of A2 posterior
to labrum; base of Mn outside of labral-
paragnathal envelope. Base of Mxp lateral
to base of Mx2; base of Mx1 lateral and
posterior to base of Mx2. Rostrum, labrum,
and paragnaths as in Fig. 2D.
Ur (Fig. 1D): 4 segments; anterior seg-
ment a genital complex of Th7 and Ab1; as
viewed dorsally, genital complex symmet-
rical (Fig. 2A). Viewed laterally (Fig. 2B,
C), genital complex with a thickened ridge
laterally on right side. Right laterally and
dorsad, posterior margin of abdominal so-
mites 1—3 asymmetrical, each with attenu-
ate, finger-like process which engages sock-
et-like depression on anterior margin of fol-
lowing somite (Fig. 2E).
Al (Fig. 3): A large segmental complex
proximally (Fig. 4A) of 19 setae separated
by incomplete ventral (anterior) arthrodial
membranes into 8 groups of 3, 2, 2+1, 3,
2, 3, 1, 2 setae+aesthetascs with setae of
same group laterally displaced relative to
each other; aesthetasc of the 3rd group
short, triangular with distal sensilla. Distal
to the complex an incompletely articulating
segment with 2 setae and short, triangular
aesthetasc with distal sensilla followed by
13 completely articulating segments with
Zt hig, Qine2 pe Did sp Dole Lawton Dee wea cee
6+1 setae+aesthetascs.
A2 (Fig. 4B, C): coxa and basis without
setae. Re 8 articulating segments with 0, 1,
VOLUME 113, NUMBER 1
79
Fig. 1.
Griceus buskeyi new genus, new species. A, Oral area, ventral, showing location of only one of
antenna | (1), antenna 2 (2), mandiblar gnathobase (3), maxilla 1 (4), maxilla 2 (5), maxilliped, obscuring
mandibular palp (6); both swimming legs | (7); B, same, showing location of origin of antenna | (1), antenna
2 (2), mandible (3), maxilla | (4), maxilla 2 (5), maxilliped (6); C, animal, left lateral; D, Th5—6 and urosome,
dorsal; E, leg 5. All scale lines are 0.01 mm.
1, 1, 1, 1, 0, 3 setae. Ri 2-segmented with
1 and 8 (4 terminal, 4 subterminal) setae.
Mn (Fig. 4D): coxa elongate with lobe at
mid-length and attenuations medially; basis
unarmed. Re 5-segmented with 1, 1, 1, 1, 1
setae. Ri apparently a lobe on the basis me-
dial to the exopod.
Mx1 (Fig. 4E): Praecoxa, coxa and basis
with poorly-developed, unarmed lobes. Ri
not distinguished. Re 1l-segmented with 1
medial and 3 terminal setae.
Mx2 (Fig. 4F): Proximal and distal prae-
coxal endites poorly-developed, unarmed;
no arthrodial membrane separating prae-
coxa and coxa. Proximal coxal endite with
2 setae, distal coxal endite with 2 setae. En-
dites of basis indistinct with 1 and 2 setae.
Re unsegmented with 7 setae.
80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 2.
=
Griceus buskeyi new genus, new species. A, Genital complex, dorsal; B, Genital complex, left lateral,
C, Genital complex, right lateral; D, Mouth area, ventral, showing location of rostrum (1), labrum (2), paragnath
(3), tip of mandibular gnathobase (4) and origin of antenna 2 (5); E, urosome, dorsal and slightly lateral, with
attenuate finger-like processes and socket-like depressions darkened. All scale lines are 0.01 mm.
Mxp (Fig. 5A): syncoxa with 1 seta but
without distinct lobes. Basis with 3 setae (2
on a distal medial lobe which is poorly de-
veloped and weakly sclerotized) and ante-
rior denticles not organized into a clear pat-
tern. Ri 4-segmented proximal to distal with
1, 2, 3, 4 setae (4, 3, 1, 2 by segmental age).
P1 (Fig. 5B): coxa with medial seta. Ba-
sis with medial denticles; medial seta
curved only toward its tip with setules
along the proximal straight part; lateral seta
absent. Re apparently 3-segmented with
distal segment missing; proximal segment
with 2 (medial and lateral) setae; middle
segment with 2 (medial and lateral) setae.
Ri 3-segmented; proximal segment with
medial seta, a pore on anterio-distal margin
(with the inner seta of the basis comprising
VOLUME 113, NUMBER 1
Fig. 3.
Von Vaupel Klein’s organ) and a lateral at-
tenuation; middle segment with 2 medial
setae; distal segment with | medial, 2 ter-
minal and 2 lateral setae.
P2 (Fig. 5C): coxa with medial seta. Ba-
sis unarmed. Re apparently 3-segmented
with distal segment missing; proximal seg-
ment with 2 (medial and lateral) setae; mid-
dle segment with 2 (medial and lateral) se-
tae. Ri 3-segmented; proximal segment
with medial seta and middle segment with
2 medial setae; distal segment with 3 me-
dial, 2 terminal and 2 lateral setae.
P3 (Fig. 5D): coxa with medial seta. Ba-
sis unarmed. Re apparently 3-segmented
with distal segment missing; proximal seg-
ment with 2 (medial and lateral) setae; mid-
dle segment with 2 (medial and lateral) se-
tae. Ri apparently 3-segmented; proximal
segment with medial seta and middle seg-
ment with 2 medial setae; distal segment
missing.
P4 (Fig. 5E): coxa with medial seta. Ba-
sis with lateral seta. Re apparently 3-seg-
mented with distal segment missing; prox-
imal segment with 2 (medial and lateral)
setae; middle segment with 1 medial seta
(lateral seta apparently missing). Ri appar-
ently 3-segmented; proximal segment with
a medial seta and middle segment with 2
medial setae; distal segment missing.
Leg 5 (Fig. 1E): coupler uniting limbs
which each bear a terminal seta.
Griceus buskeyi new genus, new species. A,
81
antenna |, left. Scale line is 0.01 mm.
CR (Fig. 1D): 4 thick apical setae; dorsal
seta small and thin.
Male.—unknown.
Etymology.—The name recognizes Ed-
ward Buskey for collecting the sample
which contained this copepod. The specific
epithet is a noun in the genitive singular.
Remarks.—The unknown, but possibly
extensive, period of time this specimen may
have remained unfixed in the collecting net
makes difficult the determination of several
possible synapomorphies involving setae or
spinules. For example, a rostrum without
spinules (filaments), antenna 2 basis with-
out setae, antenna 2 endopod with four ter-
minal setae and mandibular exopod with
one seta on the terminal segment all rep-
resent potential apomorphies that should be
verified when better preserved specimens
are available. For this reason, we have cho-
sen to emphasize the apparently derived na-
ture of the following changes in shape of
somites and appendage segments which are
presumed apomorphies for species of Gri-
ceus: interlocking extensions of abdominal
somites dorsally; antenna 2, mandible and
maxilla 1 originating posterior to the la-
brum and paragnaths; coxal endite of the
mandible lying outside of the labral parag-
nathal envelope; praecoxal and coxal en-
dites of maxilla 1 unarmed; praecoxal en-
dites of maxilla 2 unarmed; endopod of
maxilliped with four articulating segments.
82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Griceus buskeyi new genus, new species. A, proximal segmental complex plus following incompletely
articulating segment of left antenna 1, ventral view, distal is down, wavy line cutoff on broken seta, circle with
X is setal scar, broken lines are incomplete arthrodial membranes (scale line as for B); B, exopod of right antenna
2 with, segments |, 2—4 and 9 indicated; C, left antenna 2, with exopod broken; D, left mandible; E, left maxilla
1; E left maxilla 2. All scale lines are 0.01 mm.
The effect of the changes in location of
the cephalic appendages may be general-
ized as follows: A2, Mn and Mx1 retain
their ancestral positions relative to one an-
other but as a group they have been drawn
back posteriorly and laterally from the usu-
al calanoid locations. The penultimate seg-
ment of the maxilliped may be a complex
VOLUME 113, NUMBER 1
Fig: 5.
83
Griceus buskeyi new genus, new species. A, left maxilliped; B, swimming leg 1, anterior; C, swim-
ming leg 2, posterior; D, swimming leg 3, posterior; E, swimming leg 4, posterior. All scale lines are 0.01 mm.
of 2nd and 5th segments with the arthrodial
membrane missing because it bears more
setae than the two segments proximal to it
(see Ferrari 1995, Ferrari & Dahms 1998).
The endopod of the maxilliped of derived
centropagoideans is poorly sclerotized and
without arthrodial membranes; setal addi-
tions during development of Acartia tonsa
Dana, 1849, Tortanus dextrilobatus Chen &
Zhang, 1965, and Epilabidocera longipe-
data (Sato 1913) suggest these endopods
are 4-segmented with one seta on proximal,
antepenultimate and penultimate segments,
and two setae on the distal segment. The
endopod of G. buskeyi is well-sclerotized;
there is one seta on the proximal, two setae
on the antepenultimate, three setae on the
penultimate and four setae on the distal seg-
ments suggesting that this 4-segmented ra-
mus is not convergent with the centropa-
goideans. The ramus of maxilla | is inter-
preted as an exopod because its quadrate
morphology and terminal crown of setae is
similar to the exopod of many heterorhab-
dids and augaptilids. We know of no cal-
anoid copepod with a maxilla 1 in which
84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
the endopod is the only ramus. Segmental
homologies of leg 5 are based on the lo-
cation of the terminal seta toward the lateral
side of the appendage. This seta is inter-
preted as the terminal seta of the distal ex-
opodal segment; the proximal segment is a
fused coxa plus basis.
We have reconsidered what kinds of in-
formation the morphology of the exopod of
A2 provides about the superfamily of Gri-
ceus. We do not follow the hypothesis of
Park (1986) that the exopod of antenna 2
of species of Augaptiloidea have only nine
segments, none of which are complexes,
while among the remaining superfamilies
there are up to 10 segments, and except for
the Eucalanoidea with the 2nd always fused
to the 3rd and 9th always fused to the 10th.
Which of the ten segments present in the
remaining superfamilies fails to form in
Augaptiloidea is not indicated by Park
(1986). Here, homologies of the exopod are
derived from the following assumption: all
segments, with the exception of the distal
segment, are serial homologues bearing at
most one medial seta. The female of Rhin-
calanus gigas Brady, 1883 has nine medial
setae, each proximal to an arthrodial mem-
brane, and a distal segment with a terminal
and two sequentially arranged medial setae
(Fig. 6A). An incomplete, distal arthrodial
membrane extends in part along the anterior
and posterior surfaces between the second
and third medial setae, while an incomplete,
distal arthrodial membrane fails to extend
to the lateral surface separating the 3rd and
4th medial setae. Giesbrecht (1892: plate
11, figs. 16, 17) shows the exopod of Eu-
calanus attenuatus (Dana 1849) and E.
crassus Giesbrecht, 1888 which can be in-
terpreted as 11-segmented, a proximal com-
plex of four segments represented by four
medial setae, followed by six articulating
segments, each with a seta, and a distal seg-
ment with a crown of three setae. We have
been able to verify only a crown of three
setae and nine medial setae each with at
least part of a distal arthrodial membrane
for these species.
Males of Temora longicornis (Miiller,
1785) have a more common calanoid mor-
phology (Fig. 6B). An elongate segment
distally with a medial seta near its mid-
length is assumed to correspond to the pen-
ultimate segment which is elongate distad
from the seta in this species. The distal seg-
ment has a crown of three setae correspond-
ing to the distal segment of R. gigas. The
2nd articulating segment with three medial
setae is assumed to be a complex of the
2nd, 3rd, and 4th segments in which the
incomplete arthrodial membranes of R. gi-
gas fail to form on T. longicornis. The five
remaining articulating segments with a me-
dial setae and distal arthrodial membrane
are the Ist and 5th—8th of R. gigas. There
are many calanoids in which the distal ar-
throdial membrane of three or more proxi-
mal segments fails to form; in these cases,
a medial setae is assumed to represent the
location of each segment. For example, in
Calanus finmarchicus (Gunnerus, 1765) the
elongate second segment shows no trace of
an arthrodial membrane on its dorsal sur-
face but four medial setae suggest this is a
complex of the 2nd—5th segments; the re-
maining distal segments, each with a medial
seta, articulate distally (Fig. 6C).
In contrast, in augaptiloidean species like
Phyllopus bidentatus Brady, 1883 and Het-
erorhabdus spinifrons (Claus 1863), a me-
dial seta on each of a set of proximal seg-
ments fails to form while the distal arthro-
dial membrane of each of those segments is
present (Giesbrecht 1892: plate 18, fig. 29
and plate 20, fig. 9). Ohtsuka et al. (1994)
describe arietellids with an elongate, prox-
imal segmental complex in which both the
seta and distal arthrodial membrane of a set
of segments apparently fail to form. Gri-
ceus buskeyi has its 1st segment with a dis-
tal arthrodial membrane but no medial seta;
the following segmental complex is inter-
preted as composed of the 2nd and 3rd seg-
ments which have neither an arthrodial
membrane nor a medial seta, plus the 4th
segment with a medial seta and a distal ar-
throdial membrane. The next four articulat-
VOLUME 113, NUMBER 1
85
Fig. 6. Rhincalanus gigas antenna 2 exopod; Temora longicornis antenna 2 exopod; Calanus finmarchicus
antenna 2 exopod. Presumed segments are numbered proximal to distal; scale lines for A and C are 0.01 mm;
B is 0.05 mm.
ing segments each with a seta are the 5th—
8th segments. The following elongate seg-
ment is an unarmed 9th segment and the
10th segment has a crown of three setae.
Our interpretation assumes that all seg-
ments homologous to those of R. gigas are
present in these calanoids. The alternate hy-
pothesis of Park (1986) that one of the ten
segments fails to form, has not been con-
sidered because we are unsure of the loca-
tion on the antennal exopod that new seg-
ments are patterned, and whether there is a
single location for segment patterning, like
the copepod maxilliped (Ferrari & Dahms
1998), or more than one location, like the
calanoid antenna | (Ferrari & Benforado
1998).
Loss of segments on some of the swim-
ming legs complicates the assignment of
this specimen to the correct calanoid family.
However, two character states suggest that
G. buskeyi belongs to the superfamily Au-
gaptiloidea. Two setae on middle segment
of endopod of swimming legs 3 and 4,
probably an ancestral calanoid state, are
present only on Epacteriscioidea, Pseudo-
cyclopoidea, Augaptiloidea, Centropago-
idea, Megacalanoidea, and some Bathypon-
tioidea (Andronov 1974, Suarez-Morales &
Iliffe 1996). We note in passing a misprint
in Table 1 of Suarez-Morales & Iliffe
(1996) in the number of inner setae on the
terminal segment of swimming legs 3 and
4 (column E); Pseudocalanoidea (Clauso-
calanoidea) are unique in having four setae
while the Spinocalanoidea with five setae
are identical to the remaining superfamilies.
The Arietellidae and Heterorhabdidae are
the only families among the above six su-
perfamilies with species in which the en-
dopod of maxilla 1 does not develop (Oht-
suka et al. 1994, Park 2000), so an assign-
86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ment to the Augaptiloidea seems reason-
able.
Among the eight families and 39 genera
of Augaptiloidea, a set of derived characters
states have been proposed only for the fam-
ily Heterorhabdidae and its genera (Park
2000), so placement of G. buskeyi in a fam-
ily is difficult. With respect to the affinity
of G. buskeyi, a small unarmed mandibular
endopod and maxilla 1 without an endopod
are character states shared only with Par-
augaptilus Wolfenden, 1904 (Arietellidae)
and a few species of Augaptilus Giesbrecht,
1889 (Augaptilidae). The absence of setae
on the praecoxal endites of maxilla 2 sug-
gests a relationship with Paraugaptilus
which has only one seta on each praecoxal
endite. All species of Augaptilus have two
setae on these endites; this is a widespread
and presumably ancestral state for the Au-
gaptiloidea. Given our limited knowledge
the Augaptiloidea and its families, the
shared similarities of G. buskeyi to species
of Paraugaptilus provides a reasonable jus-
tification for placing this new genus in the
Arietellidae.
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Andronovy, V. N. 1974. Filogeneticheskie otnosheniya
krypnykh taksonov podotryada Calanoida
(Crustacea, Copepoda).—Zoologicheskii Zhyr-
nal 53:1002—1012. [Phylogenetic relationships
of the large taxa within the suborder Calanoida
(Crustacea, Copepoda).—translated, 1980, by
Alahram Center for Scientific Translations]
. 1991. On renaming of some taxa in Calanoida
(Crustacea).—Zoologicheskii Zhyrnal 70:133-
134 [in Russian, English summary].
Bowman, T. E. 1987. Comment on the proposed pre-
cedence of Pseudocalanidae Sars, 1901 (Crus-
tacea, Copepoda) over Clausocalanidae Gies-
brecht, 1892.—Bulletin of Zoological Nomen-
clature 44:129.
Brady, G. S. 1883. Report on the Copepoda. Part 23.—
Report on the Scientific Results of the voyage
of H.M.S. “‘Challenger’’ during the years 1873-—
1876, Zoology 8:1—142, 55 pls.
Claus, C. 1863. Die freilebenden Copepoden mit be-
sonderer Beriicksichtigung der Fauna Deutsch-
lands, der Nordsee und des Mittelmeeres. W.
Engelmann, Leipzig, 230 pp., 37 pls.
Chen, Q., & S. Zhang. 1965. The planktonic copepods
of the Yellow Sea and the East China Sea I.
Calanoida.—Studia Marina Sinica 7:20—-131 +
53 pls.
Dana, J. D. 1849. Conspectus crustaceorum, quae in
orbis terrarum circumnavigatione, Caroli
Wilkes, e classe Reipublicae foederatae duce,
lexit et descripsit Jacobus D. Dana.—Ameri-
can Journal of Science (2) 8:276—285, 424-
428.
Ferrari, F D. 1995. Six copepodid stages of Ridge-
wayia klausruetzleri, a new species of copepod
crustacean (Ridgewayliidae, Calanoida) from
the barrier reef in Belize, with comments on
appendage development.—Proceedings of the
Biological Society of Washington 108:180—
200.
, & A. Benforado. 1998. Setation and setal
groups on antenna | of Ridgewayia klausruet-
zleri, Pleuaromamma xiphias, and Pseudocalan-
us elongatus (Crustacea: Copepoda: Calanoida)
during the copepodid phase of their develop-
ment.—Proceedings of the Biological Society
of Washington 111:209-—221.
, & H.-E. Dahms. 1998. Segmental homologies
of the maxilliped of some copepods as inferred
by comparing setal numbers during copepodid
development.—Journal of Crustacean Biology
18:298-307.
Giesbrecht, W. 1888. Elenco dei Copepodi pelagici
raccolti dal Tenete di vascello Gaetano Chier-
chia durante il viaggio della R. Corvetta “‘Vettor
Pisani’? negli anni 1882—1885 e dal Tenete de
vascello Francesco Orsini nel Mar Rosso, nel
1884.—Rendiconti della Reale Accademia dei
Lincei Classe di Scienze fisiche, matematiche e
naturali 4, semestre 2:284—287, 330-338.
. 1889. Elenco dei Copepodi pelagici raccolti
dal Tenete di vascello Gaetano Chierchia dur-
ante il viaggio della R. Corvetta “‘Vettor Pisa-
ni” negli anni 1882—1885 e dal Tenete de vas-
cello Francesco Orsini nel Mar Rosso, nel
1884.—Rendiconti della Reale Accademia dei
Lincei Classe di Scienze fisiche, matematiche e
naturali 5, semestre 1:811—815, semestre 2:24—
20:
. 1892. Systematik und Faunistik der pelagisch-
en Copepoden des Golfes von Neapel und der
angrenzenden Meeres-abschnitte.—Fauna und
Flora des Golfes von Neapel und der angren-
zenden Meeres-abschnitte 19:1—831 + 54 pls.
Gunnerus, J. E. 1770. Nogle smaa rare og meesten-
deelen nye Norske Sgedyr.—Skrifter, som udi
det Kigbenhavnske Selskab af Laerdoms og Vi-
denskabers Elskere, 1765-1769, 10:166—176.
International Code of Zoological Nomenclature. 1985.
Articles 62—64. P. 119 in W. D. L. Ride et al.,
eds., 3rd Edition. University of California Press,
Berkeley, 338 pp.
VOLUME 113, NUMBER 1
International Commission on Zoological Nomencla-
ture 1988. Opinion 1503: Pseudocalanidae Sars,
1901 (Crustacea, Copepoda) not to be given
precedence over Clausocalanidae Giesbrecht
1892.—Bulletin of Zoological Nomenclature
43:228-229.
Miiller, O. EK 1785. Entomostraca seu Insecta Testacea
quae in aquis Daniae et Norvegicae reperit, des-
cripsit et iconibus illustravit Otho Fridericus
Miiller. EF W. Thiele, Lipsiae & Havniae, 134
pp. + 21 pls.
Ohtsuka, S., G. A. Boxshall, & H. S. J. Roe. 1994.
Phylogenetic relationships between §arietellid
genera (Copepoda: Calanoida), with the estab-
lishment of three new genera.—Bulletin of the
Natural History Museum London (Zoology) 60:
105-172.
87
Park, T. 1986. Phylogeny of calanoid copepods.—Syl-
logeus 58:191—196.
. 2000. Taxonomy and distribution of the ma-
rine calanoid copepod family Heterorhabdi-
dae.—Scripps Institution of Oceanography Bul-
letin 31 (in press).
Sato, C. 1913. Fuyusei-Tokyakurui.—Suisan Chosa
Hokoku 1:28-—29.
Suarez-Morales, E., & T. M. Iliffe. 1996. New super-
family of Calanoida (Copepoda) from an an-
chialine cave in the Bahamas.—Journal of
Crustacean Biology 16:754—762.
Wolfenden, R. N. 1904. Notes on the Copepod of the
North Atlantic Sea and the Farée Channel.—
Journal of the Marine Biological Association of
the United Kingdom (new series) 7:110—146,
pee.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):88—-94. 2000.
Studies on the Crustacea of the Turks and Caicos Islands, British
West Indies. IV. Heteromysis (Heteromysis) spottet, a new species
(Peracarida: Mysidacea: Mysidae) from Pine Cay
W. Wayne Price and Richard W. Heard
(WWP) Department of Biology, University of Tampa, Tampa, Florida 33606, U.S.A.;
(RWH) University of Southern Mississippi, Institute of Marine Sciences,
Ocean Springs, Mississippi 39566-7000, U.S.A.
Abstract.—During a survey of the marine Crustacea in the vicinity of Pine
Cay, Turks and Caicos Islands, an undescribed mysid, Heteromysis spottei, new
species, was collected at depths ranging from 1 to 18 m. The new species
belongs to the subgenus Heteromysis Bacescu and can be distinguished from
the five other members of that subgenus known from the western Atlantic by
the setation of the telson. In Heteromysis spottei the telson has spine-setae
along the entire length of the lateral margins, spinules along the entire margins
of the cleft, and a pair of spine-setae on each apical lobe, the outer being nearly
three times longer than the inner. A key and diagnostic table to the species of
the subgenus Heteromysis from the Northwest Atlantic is presented.
Excluding the commensal palaemonid
shrimps, which have been addressed in a
separate series of publications (Heard &
Spotte 1991, Heard et al. 1993, Spotte et al.
1994, Spotte & Bubucis 1996, Heard &
Spotte 1997), this is the fourth contribution
in a series on the crustacean fauna of the
Turks and Caicos Islands. The first three re-
ports dealt with marine isopods (Kensley &
Heard 1991, Schotte & Heard 1991, Schotte
et als 199):
This report presents the description of a
new species of mysid belonging to the ge-
nus Heteromysis Smith, 1873. The new spe-
cies, which is referred to the subgenus Het-
eromysis (Heteromysis) Smith 1873 sensu
Bacescu, 1968, was collected in water
depths ranging from 1 to 18 m in the vicin-
ity of Pine Cay.
The type material was deposited in the
National Museum of Natural History
(USNM), Smithsonian Institution, and the
Gulf Coast Research Laboratory (GCRL).
We follow the setal classification of Watling
(1989) in which a seta is defined as an ar-
ticulated cuticular outgrowth of the integu-
ment. The term “‘spine-seta’’ as used here
refers to any seta with spine-like character-
istics; a spine is defined as a non-articulated
extension of the cuticle.
Heteromysis (Heteromysis) spottei, new
species
Figs. 1-2
Material examined (all material from
Turks and Caicos).—Holotype: adult @
(Length ,[L] 3:1 mm), JUSNM) 282712:
fringing reef off Pine Cay, live bottom/
sand, depth 18 m, J. A. McLelland (coll.),
suction device, 12 Nov 1989. Paratypes:
adult ¢ (L3.5mm), USNM 282720; adult
6, damaged, GCRL 1346, same collection
data as holotype.—1 adult ¢d, 2 29, (all
damaged) Pine Cay, shallow inner reef,
sand-coral rubble substratum with associ-
ated sponges and cnidarians, depth 4 m, 1
mm mesh dredge net, R. Heard, J. Mc-
Lelland, P. Bubucis, & S. Spotte (colls.), 5
Nov 1988.—16d6 (damaged), Pine Cay
(Rock-a-Wash Cay), depth 1 m, Neogoniol-
ithon & sponge washings, R. Heard (coll.),
VOLUME 113, NUMBER 1
30 Oct 1988.—2¢ 6, 1 ¢ (all damaged),
same data as holotype.
Diagnosis.—Article 3 of antennular pe-
duncle without distomedial flagellated
spine-seta; thoracic endopod 3 with 3 flag-
ellated spine-setae on medial margin of
merus, medial margin of carpo-propodus
without flagellated spine-setae; pleopods
uniramous, reduced to simple setose plates
with no modified spine-setae in either sex;
endopod of uropod armed with 1 small
spine-seta near statocyst; lateral margins of
telson armed along entire length with 10—
12 spine-setae per margin (including apical
spine-setae), posterior-most lateral spine-
seta 1.6—-1.7 times length of preceding
spine-seta, extending beyond posterior end
of telson; outer apical spine-seta nearly 3
times longer than inner; cleft completely
armed with 18—20 spinules.
Description.—General body form (Fig.
1A): moderately robust; carapace with an-
terior margin produced into pointed trian-
gular rostrum; posterior dorsal margin
emarginate, partly exposing thoracic seg-
ment 8; anterolateral lobes rounded.
Antennule peduncle (Fig. 1B): article 1
slightly shorter than article 3, with 3 plu-
mose and 2 simple spine-setae on distolat-
eral process, dorsomedial longitudinal ridge
with 4 strong simple distal spine-setae; ar-
ticle 2 compressed with 2 distomedial
spine-setae, | simple and 3 plumose spine-
setae near middle of distal margin; article 3
with simple spine-seta on medial margin, 3
simple distomedial spine-setae, 3 plumose
spine-setae near distolateral margin, | sim-
ple spine-seta on dorsolateral surface, lack-
ing flagellated spine-seta, males with small
moderately setose lobe on ventral surface.
Antenna (Fig. 1C): scale slightly shorter
than peduncle, 3.0—3.2 times as long as
maximum width, medial margin strongly
convex, lateral margin straight, all margins
setose, lacking distal article; antennal pe-
duncle having 3 articles; article 1 incon-
spicuous; article 2 just over 1.5 times lon-
ger than article 3, short plumose spine-seta
near distolateral border, 2 simple and 2 plu-
89
mose distomedial spine-setae; article 3 with
1 plumose and 3 simple distomedial spine-
setae, 3 plumose spine-setae along lateral
margin.
Eyes (Fig. 1A): large, oval, directed lat-
erally, distal part of eye stalk wider than
cornea, lacking ocular tooth; cornea large,
oval.
Mandibles (Fig. 1D—F): molar, incisor,
and lacina mobilis as illustrated. Palp 3-seg-
mented; article | small, inconspicuous; ar-
ticle 2 expanded, medial margin with 5—10
simple spine-setae, lateral margin with 4—5
proximal plumose spine-setae and | simple
distolateral spine-seta; article 3 half as long
as 2, medial margin with | plumose spine-
seta, distal part armed with | simple and 9—
10 barbed spine-setae, 1 simple spine-seta
on medial surface.
Labrum and paragnaths (Fig. 1D, G): as
illustrated.
Maxillule (Fig. 1H): outer lobe with 11
stout apical and 3 subapical spine-setae; in-
ner lobe with 3 long, distally curved, serrate
spine-setae, 2 plumose and 2 simple spine-
setae distally, and 1 plumose and | simple
spine-seta on distomedial margin.
Maxilla (Fig. 11): as illustrated; exopod
with 13—15 plumose spine-setae on disto-
lateral margin.
Thoracic endopods 1-8 (Fig. 2A-—G):
thoracic endopods 1 and 2 as illustrated.
Thoracic endopod 3, merus approximately
1.6 length of ischium and equal in length
to carpo-propodus, medial margin with 3
flagellated and 3 simple spine-setae, lateral
margin with | distal simple spine seta; me-
dial margin of carpo-propodus with 4—5
simple spine-setae, 2 simple and 2 stout ser-
rate spine-setae on distomedial margin; dac-
tyl small, with long, slightly curved claw
on distal end surrounded by several simple
spine-setae. Thoracic endopod 4, merus
about 1.5 length of ischium; carpo-propo-
dus about %4 length of merus, with 3 articles
(distal 2 subequal, combined length slightly
longer than proximal article); dactyl mi-
nute, papillate, lacking terminal claw. Tho-
racic endopod 5, ischium and merus sub-
90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
=
Sy
Zp ~
Heteromysis spottei, new species. A, adult 2, dorsal view; B, antennular peduncle; C, antennal
peduncle and scale; D, labrium & right mandibular palp; E, KE right mandible, inner and upper aspects, respec-
tively; G, paragnaths; H, maxillule; I, maxilla. Scale 1, A = 0.5 mm; C = 0.2 mm; B, D-E I = 0.1 mm; A =
0.05 mm; Scale 2, G = 0.1 mm.
equal in length; carpo-propodus with 4 ar-
ticles, distal 3 subequal in length, each half
as long as proximal article, ultimate article
with 6—7 simple spine-setae (2 sickle-
shaped with strong articulated bases); dac-
tyl small, papillate with long distal, slender
claw. Thoracic endopod 6, carpo-propodus,
ischium and merus subequal in length; car-
po-propodus with 5 articles, distal 4 sub-
equal in length, each about % as long as
proximal article; dactyl small, papillate with
distal serrate slender claw. Thoracic endo-
pod limb 7, ischium slightly longer than
merus, merus slightly longer than carpo-
propodus; carpo-propodus with 3 articles,
proximal distinctly longer than each of dis-
tal articles; dactyl small, papillate with dis-
tal serrate slender claw. Thoracic endopod
8 (1 damaged limb available for study),
long, attenuated, at least 4% longer than other
thoracic endopods; carpo-propodus with at
least 5 articles.
Thoracic exopods: exopod 1 with 8 arti-
cles; exopods 2—8 with 9 articles.
Thoracic sternal processes: median spi-
niform processes on sterna 3—7 in males.
VOLUME 113, NUMBER 1
9]
Fig, 2.
Heteromysis spottei, new species. A-—G endopods of thoracic limbs 1—7; H, telson; I, uropod. Scale
1, H = 0.1 mm; Scale 2, A = 0.1 mm, B—G = 0.2 mm; Scale 3, I = 0.2 mm.
Pleopods: without sexual dimorphism;
reduced to uniramous plates with no mod-
ified spine-setae, similar in form.
Uropods (Fig. 21): exopod about 1.2
times longer than endopod, lateral margin
straight, medial margin slightly convex, all
margins setose; endopod linguiform with 1
spine-seta on medial margin in region of
statocyst, all margins setose.
Telson (Fig. 2H): 0.8 times length of ex-
opod of uropod, 1.4—1.5 times as long as
maximum width, lateral margins slightly
concave, armed along entire length with 10—
12 spine-setae per margin (apical spine-setae
included), increasing in length posteriorly,
most posterio-lateral spine-seta 1.6—1.7
times length of preceding spine-seta, extend-
ing beyond posterior lobes of telson; outer
apical spine-seta 2.8—3.0 times longer than
inner; cleft, depth 0.25 length of telson,
completely armed with 18—20 small spines.
Etymology.—This species is named for
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Neen nner nnn rr nn reese SSS
Cenvic ERS OG EC OiGaL Cc OVr-8e [eos [BUUDIUR JO ORI YIPIM:yIsUsT
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ouou ouou é ouou ouou ouou spodogjd ayeu poyipoyy
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-ug[s /-9 ‘ajduiis = ‘ropusys 6 ‘ajduris = ‘tapusys / ‘aydurts ouou = -UaTS 6-8 ‘ayduuts ‘Jopus[s ‘ojduts = -1v9 JO UTSIeU [eIpoW UO deIaS-9UIdS
ajyounpod
juasqe juasqe juosqe juasqe juasqe juasqe Jejnuusjue uo vjas-oulds payey[aseyy
canner nr SSS Ss
1ajJods inuvasainu 1jaanou DUDIIXIU psowusof suds] Jquoevieyo
See
sa1oadg
anne nnn nn SSS
“ONUL[IY UI9ISOMYIOU IY) Ul SISdMOsLIJAF] SNUISQNS dy} JO S1d}dvIeYO J[Npe oytoods puke uoWWIOD—'| 2IqQR I],
VOLUME 113, NUMBER 1
Stephen Spotte in recognition of his many
contributions to marine science.
Habitat.—Heteromysis spottei was col-
lected from habitats having a variety of cal-
careous algae, sponges, anthozoans, and
other sessile forms. It occurred in depths of
less than 1 m on the Caicos Banks, 3—4 m
in the back reef area, and 18 m on the outer
fringing reef. Unfortunately, our collecting
techniques were too general to determine if
the species was associated with a specific
sessile host.
Distribution.—This species is presently
known only from waters immediately ad-
jacent to Pine Cay, Turks and Caicos Is-
lands, British West Indies.
Remarks.—The dichotomous key and Ta-
ble 1 present some of the salient characters
that distinguish these western Atlantic spe-
cies of the subgenus Heteromysis.
Key to western Atlantic species of the
subgenus Heteromysis Smith, 1873 sensu
Bacescu, 1968
(species lacking flagellated spine on
segment 3 of antennular peduncle and
sexually dimorphic pleopods).
1. Endopod of uropod with 12 or more spines
along medial margin from region of statocyst
1) EL HAIRS ARS rd ee ara Z
—Endopod of uropod with no spines or one
spine medially, near statocyst ......... =
2. Margins of telsonic cleft with spinules along
entire length; antennal scale 3 times or less
as long as maximum width .......... 3
—Margins of telsonic cleft with spinules
only in apical part; antennal scale at least 3.5
times as long as maximum width.........
BNA 6103s Heteromysis elegans Brattegard, 1974
3. Carpo-propodus of third thoracic endopod
with 3 pairs of stout spine-setae on distome-
SCS WiLL Ee ee Re Re ce na a
.... Heteromysis formosa S. 1. Smith, 1873
—Carpo-propodus of third thoracic endo-
pod with no stout spine-setae on medial
MURS PMN Rese ctl a Nag Mine es Oe A gers 'y. gs
Heteromysis mexicana Escobar-Briones &
Soto, 1990
4. Endopod of uropod with no spine-setae me-
dially, near statocyst
Aas Ouse ee ere le,)ew te =, sw We fa, ee
93
....Heteromysis mureseanui Bacescu, 1986
—Endopod of uropod with | medial spine-
Sever ar SIMLOCYSE oo. vee ke ee ees 5
5. Posterior half of lateral margins of telson
armed with spine-setae; margins of telsonal
cleft with spinules in apical half only .....
hy ast Heteromysis nouveli Brattegard, 1969
—Lateral margins of telson armed with
spine-setae along entire length; margins of
telsonal cleft with spinules along entire
length
eis Wan ee eke 'e Shia” Gee bi Wen. fe ete) We “EOS eee ts se fw
Heteromysis spottei appears most similar to
H. nouveli, but differs in several aspects, espe-
cially in the setation of the telson and third tho-
racic endopod (see Key, Table 1). Except for H.
(Heteromysis) waitei W. Tattersall, 1927, a
South Australian species, the distinctive telson
setation of H. spottei differs from all other de-
scribed species of the subgenus Heteromysis.
Several characters distinguish H. spottei from H.
waitei including the presence of a single, instead
of 3—4, spine-setae adjacent to the statocyst on
the uropodal endopod and the absence of ocular
teeth (see W. Tattersall 1927).
Acknowledgments
Support for the collection of specimens
used in this study was sponsored by the
Oakleigh L. Thorne Foundation through a
grant to Stephen Spotte. We thank Oakleigh
B. Thorne, members and employees of the
Meridian Club, and the Turks and Caicos
government for support and encourage-
ment. We are grateful to Steve Spotte for
his help, interest, and overall coordination
of the field work for the faunal survey. Pa-
tricia Bubucis, Jerry McLelland, and Steve
Spotte assisted in the collection of speci-
mens and provided other helpful assistance.
The first author was supported through a
University of Tampa Faculty Development
Grant.
Literature Cited
Bacescu, M. 1968. Heteromysini nouveaux des eaux
Cubaines: trois especes nouvelles de Hetero-
mysis et Heteromysoides spongicola n.g.,
n.sp.——Revue Roumaine de Biologie, Serie de
Zoologie 13:221-—237.
94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
. 1986. Heteromysis mureseanui n.sp. and Kal-
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OTs
Brattegard, T. 1969. Marine biological investigations
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Escobar-Briones, E., & L. A. Soto. 1990. Heteromysis
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Crustacea of the Turks and Caicos Islands, Brit-
ish West Indies. HI. Records of marine Isopoda
from Pine Cay, Ft. George Cay, Water Cay, and
adjacent waters.—Gulf Research Reports 8:
251-257.
Smith, S. I. 1873. Systematic catalogue of the inver-
tebrates of the southern New England and ad-
jacent waters. Crustacea: Pp. 545-580 in A. E.
Verrill, Report upon the invertebrate animals of
Vineyard Sound and the adjacent waters, with
an account of the physical characters of the re-
gion: Pp. 295-778 in S. FE Baird, Report on the
condition of the sea fisheries of the south coast
of New England in 1871 and 1872.—United
States commission of Fish and Fisheries 7:1—
852.
Spotte, S., & P. M. Bubucis. 1996. Diversity and abun-
dance of caridean shrimps associated with the
slimy sea plume (Pseudopterogorgia ameri-
cana) at Pine Cay, Turks and Caicos Islands,
British West Indies.—Marine Ecology Progress
Series 113:229-—232.
, R. W. Heard, & P. M. Bubucis. 1994. Ponto-
niine shrimps (Decapoda: Caridea: Palaemoni-
dae) of the northwest Atlantic. IV. Periclimenes
antipathophilus new species, a black coral as-
sociate from the Turks and Caicos Islands and
Eastern Honduras.—Bulletin of Marine Science
55:212—227.
Tattersall, W. M. 1927. Australian opossum shrimps
(Mysidacea).—Records of the South Australian
Museum 3:235-—257.
Watling, L. 1989. A classification system for crusta-
cean setae based on the homology concept. Pp.
15-26 in B. E. Felgenhauer, L. Watling & A.
B. Thistle, eds., Crustacean Issues 6, Functional
morphology of feeding and grooming in Crus-
tacea, A. A. Balkema, Rotterdam, 225 pp.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):95-103. 2000.
Gynodiastylis laciniacristatus, a new species (Crustacea: Cumacea)
from Australia
Sarah Gerken and Jennifer Gross
Darling Marine Center, School of Marine Sciences, University of Maine
Walpole, Maine 04573, U.S.A.
Abstract.—A new species, Gynodiastylis laciniacristatus (Crustacea: Cuma-
cea) is described from two disparate localities on the Australian shelf, the Bass
Strait (Southeast) and the Dampier Archipelago (West), from depths of 48—85
m. The ovigerous female is completely figured. No males were collected. The
new species is distinguished from all other species of Gynodiastylis by a large
blunt process on the ischium of pereopod 4 and by the pattern of toothed ridges
on the carapace. G. mutabilis Hale, 1946 and G. ornata Hale, 1946 have similar
ridge patterns, however G. laciniacristatus has a distinct sharp corner on the
anterolateral prominence of the carapace and rudimentary exopods on pereopods
3 and 4, in addition to the large blunt process on the ischium of pereopod 4.
The genus Gynodiastylis is known pri-
marily from the Southern hemisphere. Hale
(1946) described the great majority of the
species in the genus, from the environs of
Australia. A new and unusual species of
Gynodiastylis was identified from the ex-
tensive cumacean collections of the Muse-
um of Victoria and is described here, as part
of an NSF PEET (Partnerships for Enhanc-
ing Expertise in Taxonomy) project.
Methods
Samples were collected with the WHOI
epibenthic sled or a Smith-MclIntyre grab.
Drawings were prepared using a camera lu-
cida on a Wild compound microscope.
Body length was measured from the tip of
the pseudorostral lobes to the posterior bor-
der of the last somite. One ovigerous fe-
male was prepared for SEM (scanning elec-
tron microscope) photography.
Family Gynodiastylidae Stebbing 1912
Gynodiastylis Calman 1911
Gynodiastylis laciniacristatus, new species
Figs. 1-5
Type material.—Holotype (NMV J45433)
1 ovigerous female, Bass Strait, Australia,
39°49.0'S, 143°24.0’E, 56 m, 20 Nov 1981.
Paratype (NMV J45434) | ovigerous °,
Bass-> “Strait, — Australia,’ °39°49;0’S,
143°24.0'E, 56 m, 20 Nov 1981. Paratypes
(NMV J45435) 1 ovigerous 2, 1 subadult
oo Bass, Strait) “Austral! 39°49 O'S,
143°24.0’E, 56 m, 20 Nov 1981. Paratype
(NMV J45436) 1 ovigerous °, Bass Strait,
Australia, 38°38.2’S, 142°35.0’E, 59 m, 20
Nov 1981. Paratype (NMV J45437) 1 ovig-
erous 2, Bass Strait, Australia, 39°06.3’S,
142°55.6’E, 81 m, 21 Nov 1981. Paratype
(NMV J45438) 1 ovigerous ¢, Bass Strait,
Australia, 40°00.0’S, 144°20.9’E, 48 m, 22
Nov 1981. Paratype (NMV J45439) 1 ovig-
erous 2, Bass Strait, Australia, 39°00.2'S,
144°33.9'E, 74 m, 23 Nov 1981. Paratype
(NMV J45440) 1 subadult 2, Bass Strait,
Australia, 39°13:6 S; 143°55:6'E; ‘85m, 23
Nov 1981. Paratype (NMV J45441) 1 sub-
adult ¢, Western Australia, (damaged),
20°1.00’S, 117°11.00’E, 48 m, 11 Jun 1983.
Paratype (NMV J45442) 1 subadult 9°,
Western Australia, 19°38.00’S, 118°6.00’E,
49 m, 13 Jun 1983.
Diagnosis.—Carapace with depression
sweeping dorsally from anterior edge, an-
terolateral prominence produced as sharp
96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
corner, with dorsal ridge bounding promi-
nence toothed, dorsal depression bounded
by sharp lateral ridges, continuing through
dorsum of pereon 5, anterior ventral margin
toothed, antennal notch oblique. Pereopod
4 with large blunt process on ischium.
Adult females less than 4 mm.
Description.—Ovigerous female, 3.5
mm. Carapace with dorsal depression, eye-
lobe present, no lenses; pseudorostral lobes
0.5 carapace length; antenna 1 visible, ex-
tending just past pseudorostral lobes. Per-
eonites 1—3 expanded ventrally as anteriorly
directed flap, pereonites 4 and 5 directed
posteriorly dorsally (Figs. 1A, 1B, 2A).
Antennule article 1 longer than articles 2
and 3 together, bearing 1 plumose and 2
simple setae proximally and 1 simple seta
distally; distomedial margin with fine hair-
like setae; article 2 bearing 1 simple and 1
plumose setae distally, medial margin bear-
ing fine hair-like setae; articles 2 & 3 sub-
equal in length, article 3 bearing 2 simple
setae; main flagellum of 2 articles bearing
1 long annulate seta, 2 long simple setae,
and 2 short simple setae terminally; acces-
sory flagellum of 1 article with 2 terminal
setae (Fig. 3A).
Mandible navicular, with 10 lifting setae;
left side with stout lacinia mobilis, incisor
quadridentate, right side with slender laci-
nia mobilis, incisor bidentate (Fig. 3B).
Maxillule of 2 lobes, outer broad lobe
bearing double row of stout setae terminal-
ly, medial margin bearing fine hair-like se-
tae; inner lobe bearing 2 tridentate, | long,
2 short setae terminally, medial margin
bearing fine hair-like setae (Fig. 3C).
Maxilla of 3 lobes; broad lobe 4 times as
wide as narrow lobe, medial margin bearing
9 simple setae, medial-distal corner bearing
2 plumose setae, distal margin bearing
many simple setae; distal-lateral corner
bearing | forked and 1 microserrate setae;
inner narrow lobe bearing 2 microserrate
and 2 simple setae apically; outer narrow
lobe bearing 5 simple setae apically (Fig.
3D).
Maxilliped 1 basis as long as all other
articles together, produced as large disto-
medial lobe, lobe bearing several short sim-
ple setae distally, lobe bearing several plu-
mose setae, setulose on distal 0.3 only; is-
chium not present, merus half as long as
carpus, medial margin bearing fine hair-like
setae; carpus bearing 3 blade-like, 3—4 bi-
dentate, several simple setae medially, 1
long plumose seta distolaterally; propodus
as long as carpus, half as broad, bearing 2
tridentate and 2 long plumose setae distally,
margins with many fine hair-like setae; dac-
tyl half as long and half as wide as propo-
dus, bearing 4 small simple setae terminally
(Fig. 5C).
Maxilliped 2 basis as long as next 2 ar-
ticles together, bearing 3 long plumose se-
tae distally; ischium not present; merus 0.5
basis length, bearing 2 plumose setae dis-
tally; carpus slightly longer than propodus,
bearing 3 plumose setae medially; propodus
bearing 2 long plumose setae distally and 3
plumose setae medially; dactyl half length
of propodus, bearing 4 simple setae termi-
nally; endite bearing 5 stout annulate setae
and 2 simple setae (Fig. 2B).
Maxilliped 3 basis 2 times as long as
next 4 articles together, medial margin bear-
ing 8 plumose setae, lined with short hair-
like setae, single plumose seta distally, dis-
tomedial corner produced as 2 teeth, lateral
margin lined with fine hair-like setae, dis-
tolateral corner bearing 4 long plumose se-
tae; ischium 0.5 basis width, slightly longer
than merus, 1 plumose setae medially, oth-
erwise margins lined with fine hair-like se-
tae; merus 0.5 length carpus, produced as
tooth at distolateral corner and bearing 1
plumose seta, medial margin lined with fine
hair-like setae and bearing | plumose and 1
simple setae distally; carpus bearing 1 plu-
mose seta on distolateral corner, medial
margin lined with fine hair-like setae and
bearing 1 plumose and 1 simple setae; pro-
podus subequal to dactyl, bearing | simple
seta on distolateral corner, medial margin
bearing 2 simple setae; dactyl bearing 4
simple setae terminally (Fig. 3E).
Pereopod 1 basis as long as all other ar-
VOLUME 113, NUMBER 1 97
eee B
Fig. 1. Gynodiastylis laciniacristatus, new species. Ovigerous ¢ (not the same individual as Figs. 2—5); A,
habitus; B, dorsal view. Scale bars are 200 microns in length.
ticles together, posterior margin bearing fine hair-like setae, anterodistal corner bear-
many simple and 2 plumose setae, postero- ing 2 plumose setae; ischium 0.5 length of
distal corner produced as 3 teeth, anterior merus, posterodistal corner produced as
margin bearing 1 simple seta, lined with tooth; merus produced as 3 small teeth on
98
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
a
a
A
a
A
a
A
a
a
a
A
A
a
a
a
a
a
a
a
a
a
\t
Pa
)
1
wee
so
mo
viv
D earn nn null
Fig 2.
illiped 2; C, telson and uropods.
Gynodiastylis laciniacristatus, new species. Ovigerous 2 paratype NMV 45434; A, habitus; B, max-
VOLUME 113, NUMBER 1 99
A, C,D 0.2mm
B,E 0.5mm
Fig. 3. Gynodiastylis laciniacristatus, new species. Ovigerous 2 paratype NMV 45434; A, antennule; B,
mandible; C, maxillule; D, maxilla; E, maxilliped 3.
i00 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
posterior margin; carpus subequal to pro-
podus; propodus twice dactyl length, bear-
ing | long and 1 short simple setae distally,
single small simple seta proximally; dactyl
bearing 1 long and 1 short simple setae ter-
minally; exopod 0.5 length basis, basal ar-
ticle bearing 2 simple setae, produced as
small tooth distally; flagellum bearing 6
long stout plumose setae (Fig. 4A).
Pereopod 2 basis longer than all other ar-
ticles together, 3 times ischium width, mar-
gins lined with many simple setae, anterior
margin produced as multiple teeth; ischium
reduced, unarmed; merus twice carpus
length, anterior margin produced as multi-
ple teeth, bearing 2 simple setae; carpus an-
terior margin produced as 2 teeth, bearing
2 simple setae; propodus subequal to car-
pus, unarmed; dactyl slightly longer than
propodus, bearing 3 simple setae terminal-
ly; exopod subequal to basis, basal article
unarmed, flagellum bearing 4 long stout
plumose setae (Fig. 4B).
Pereopod 3 basis as long as next 2 arti-
cles together, anterior margin produced as
multiple teeth, bearing 5 plumose, 1 annu-
late and 2 annulate plumose setae, posterior
margin bearing 2 simple setae; ischium 0.2
length merus, bearing 1 simple seta; merus
twice carpus length, bearing 2 annulate and
4 simple setae; carpus twice propodus
length, bearing 6 annulate and 2 simple se-
tae distally; propodus twice dactyl length,
bearing single annulate seta distally; dactyl
bearing 2 simple setae terminally; exopod
biarticulate, rudimentary, bearing 1 simple
seta apically (Fig. 4C).
Pereopod 4 coxa bearing several stout
simple setae; basis as long as next 2 articles
together, anterior margin bearing 2 plu-
mose, 1 annulate plumose, 7 annulate, and
3 simple setae, produced as multiple teeth
distally, 1 plumose seta mid article; ischium
0.3 length merus, unarmed, produced as
large posteriorly directed blunt lobe; merus
3 times carpus length, bearing 7 annulate
setae, produced as multiple scales, posterior
margin produced as 3 teeth; carpus slightly
longer than propodus, bearing 7 annulate
setae, produced as 2 teeth proximally; pro-
podus slightly longer than dactyl, bearing 2
annulate setae; dactyl bearing single stout
seta with single setule; exopod biarticulate,
rudimentary, bearing | simple seta apically
(Fig. 5A).
Pereopod 5 basis as long as next 2 arti-
cles together, bearing 6 plumose, | annulate
plumose and | annulate setae, produced as
small blunt processes on posterior margin;
ischium 0.5 length merus, bearing | simple
seta; merus slightly longer than carpus,
bearing 5 simple and 1 annulate setae, pos-
terior margin produced as scales; carpus
bearing 7 annulate setae; propodus subequal
to carpus, bearing 1 annulate seta; dactyl
0.5 length propodus, bearing 1 stout and 1
slender setae terminally (Fig. 5B).
Telson equal in length to pleonite 6, post-
anal section negligible, unarmed (Fig. 2C).
Uropod peduncles equal in length to tel-
son, bearing single seta at distomedial cor-
ner; rami shorter than peduncles; endopod
biarticulate, article 1 bearing 1 short stout
seta at distomedial corner, article 2 bearing
2 short stout setae medially, single long seta
terminally; exopod biarticulate, article 1
half length article 2, unarmed, article 2
bearing 2 short setae laterally, single long
stout seta terminally; all setae on rami bear-
ing single thick setule apically (Fig. 2C).
Etymology.—laciniacristatus from the
Latin lacinia, meaning jagged and crista,
meaning ridge.
Remarks.—Gynodiastylis laciniacrista-
tus is distinguished from all other Gyno-
diastylis by the large blunt process on the
ischium of pereopod 4. Gynodiastylis laci-
niacristatus is superficially similar to both
G. ornata Hale, 1946 and G. mutabilis
Hale, 1946. However, there are obvious dif-
ferences in addition to the process on pe-
reopod 4. Neither G. ornata nor G. muta-
bilis have exopods on pereopods 3 and 4 of
the female, while they are present in G. la-
ciniacristatus. The uropod endopod in G.
mutabilis is uniarticulate, while in G. laci-
niacristatus the uropod endopod is biartic-
ulate. The lateral margins of the telson in
VOLUME 113, NUMBER | 10]
A,B, C 0.5mm
Fig. 4. Gynodiastylis laciniacristatus, new species. Ovigerous paratype NMV 45434; A, pereopod 1; B,
pereopod 2; C, pereopod 3.
102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
C 0.2 mm
A, B, 0.5 mm
Fig. 5. Gynodiastylis laciniacristatus, new species. Ovigerous 2 paratype NMV 45434; A, pereopod 4; B,
pereopod 5; C, maxilliped 1.
VOLUME 113, NUMBER 1
G. mutabilis are produced as at least one
pair of teeth, while in G. laciniacristatus
the telson lateral margins are entire. The
carapace of G. laciniacristatus has a dis-
tinct anterolateral prominence produced as
a sharp corner, while G. ornata has no such
prominence.
Acknowledgments
The Museum of Victoria kindly loaned
the material used in the study. The National
Science Foundation (Grant DEB95-21783,
L. Watling, I. Kornfield PI.’s) supported
this research as part of the Partnerships to
Enhance Expertise in Taxonomy (PEET)
103
program. The Gulf of Maine Foundation
supported this research as a Summer Un-
dergraduate Research Experience (SURE)
for Jennifer Gross.
Literature Cited
Calman, W. T. 1911. On new or rare Crustacea of the
Order Cumacea from the collection of the Co-
penhagen Museum, Part IJ.—Transactions of
the Zoological Society of London 18:341—399.
Hale, H. 1946. Australian Cumacea, No. 12, the Fam-
ily Diastylidae (part 2) Gynodiastylis and relat-
ed genera.—Records of the South Australian
Museum 8(3):357—444.
Stebbing, T. R. R. 1912. The sympoda, Part 6.—An-
nals of the South African Museum 10:129—176.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):104—123. 2000.
Revision of the subterranean amphipod genus Spelaeogammarus
(Bogidiellidae) from Brazil, including descriptions of three new
species and considerations of their phylogeny and biogeography
Stefan Koenemann and John R. Holsinger
Department of Biological Sciences, Old Dominion University,
Norfolk, Virginia 23529-0266, U.S.A.
Abstract.—Three new subterranean amphipods of the genus Spelaeogam-
marus da Silva Brum, 1975, are described from eastern Brazil, bringing the
total number of species in the genus to four. Based on the examination of type-
material of Spelaeogammarus bahiensis, a comparative diagnosis of all four
species of Spelaeogammarus, including the new species Spelaeogammarus spi-
nilacertus, Spelaeogammarus trajanoae, and Spelaeogammarus santanensis, 1s
given. A diagnosis for the genus is also provided. The occurrence of these
species in caves that are separated from each other in discontinuous karst areas
is biogeographically significant for the family Bogidiellidae in continental
South America.
Exploration of caves in eastern Brazil in
the early 1970s resulted in the discovery of
the new bogidiellid genus and species Spe-
laeogammarus bahiensis (da Silva Brum,
1975). The specimens were collected from
a cave near Curaca, capital of the district
Matamuté, in the state of Bahia. Subse-
quently, between 1989 and 1993, many ad-
ditional specimens from various caves in
Bahia were collected by Brazilian speleol-
ogists and sent to us for identification. Be-
cause these specimens differed morpholog-
ically from the description of Spelaeogam-
marus bahiensis, we borrowed paratypes of
this species from the Museu Nacional in
Rio de Janeiro, for a comparison with the
new material. The paratypes enabled us to
identify and describe three new species and
also to diagnose the genus Spelaeogam-
marus. In addition, a key to the four species
of the genus is provided as well as a table
detailing morphological differences.
The holotypes of the new species are de-
posited in the Museu Nacional in Rio de
Janeiro (MNRJ), Brazil, as indicated.
Genus Spelaeogammarus da Silva Brum,
1975
Spelaeogammarus da Silva Brum, 1975:
125-128.
Type species (by monotypy): Spelaeo-
gammarus bahiensis da Silva Brum, 1975.
Diagnosis.—Eyes absent. Body smooth,
unpigmented. Coxal plates 1—2 small, wider
than long; plates 3-6 longer than wide,
overlapping. Antenna 1 about 45-50% of
body length, primary flagellum longer than
peduncle, with 16—20 segments. Accessory
flagellum with 4—5 segments. Antenna 2
flagellum bearing 7—10 segments. Mandib-
ular palp 3-segmented. Maxilla 1 with sym-
metrical, 2-segmented palp; inner plate with
3 plumose setae; outer plate bearing 6—7
serrate spines. Inner plate of maxilliped
bearing apically 2 bifid (y-shaped) spines;
outer plate with 3 or 4 blade-like spines api-
cally and subapically. Propodus of gnatho-
pod 1 larger than that of gnathopod 2. Dac-
tyls of both gnathopods distinctly serrate
along inner margins. Pereopods without any
trace of lenticular organs; pereopods 5—7
105
VOLUME 113, NUMBER 1
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soulds ayes /
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106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
bases broad, propodus and/or carpus with
long, bifurcate setae. Pleopods and uropods
unmodified. Pleopods biramous, with 3-
segmented outer ramus and 1-segmented in-
ner ramus, rami subequal in length. Uro-
pods biramous: peduncle of uropod | with
3 or 4 large basiofacial (ventrolateral)
spines; uropod 3 with subequal, 1-segment-
ed rami, outer ramus bearing a row of long,
bifurcate setae along medial margin. Telson
typically longer than wide, apex with shal-
low excavation, bearing apical and subapi-
cal spines. Coxal gills present on pereopods
4—6. Oostegites linear and elongate, on pe-
reopods 2—5. No sexual dimorphism in any
characters.
Key to the species of Spelaeogammarus
(based on males and females)
1. Accessory flagellum of antenna 1 with 4 seg-
ments; flagellum of antenna 2 with 7 seg-
ments; propodus of gnathopod | slightly larg-
Cr ‘thai DasiS <a ges epee ee oe 2
Accessory flagellum of antenna 1 with 5
segments; flagellum of antenna 2 with 8—10
segments; propodus of gnathopod 1 much
larger than basis’ #yn.-. 6... .. S. santanensis
2. Coxal plate 6 bearing 1 spine, setae absent;
outer plate of maxilla 1 with 6 serrate spines
and | plumose seta; inner plate of maxilliped
with. 4; plumose.setae... . 2: .Naajun. 52 3
Coxal plate 6 bearing 1 spine and about 20
setae; outer plate of maxilla 1 with 7 serrate
spines; inner plate of maxilliped without plu-
MOSE Sta 92 4. Bon eke kh eee S. bahiensis
3. Anterior margin of basis of gnathopod 1 with
5—9 short setae; coxal plate 5 bearing 1 spine
and 17-18 setae S. trajanoae
Anterior margin of basis of gnathopod 1
with 2—4 spines (or longer setae) and 1 short
seta; coxal plate 5 bearing 1 spine and about
O SSCAAG oa aa oes Oo eae S. spinilacertus
© wr 6; © 16 @ 6 te 8, ae
Spelaeogammarus spinilacertus, new
species
Figs. 1-5, 6c
Material examined.—Estado do Bahia,
Brazil: holotype male (6.1 mm) and 1 para-
type male (7.5 mm) from Baixa do Salitre
Cave, Iraquara, collected by J. A. Cardoso,
Sep 1993; 4 paratypes (3 females, 1 male)
from Baixa do Salitre Cave, collected by L.
Mendes & E. Rubbioli, 24 May, 1997; al-
lotype female (8.1 mm) and 2 paratypes (1
male, 1 female), from Jaburu Cave, Ira-
quara, collected by J. A. Cardoso, Sep
1993"
The holotype and allotype are dissected
and mounted on microscope slides in
Faure’s medium. Holotype (MNRJ 13340)
and allotype are deposited in the Museu Na-
cional. The remaining 7 paratypes are re-
tained in the research collection of the sec-
ond author.
Diagnosis.—Spelaeogammarus spinila-
certus is easily distinguished from other
species in the genus by 2—4 spines or setae
of corresponding length on the distoanterior
margin of basis of gnathopod 1. Largest
male 10.5 mm, largest female 10 mm.
Description.—Antenna | (Fig. 2a) about
45-50% of body length. Peduncular seg-
ments 1—3 gradually decreasing in length;
peduncular segment 1 with 3—4 spines; pe-
duncular segment 2 with 2—3 short spines;
peduncular segment 3 bearing O-—2 spines.
Primary flagellum longer than peduncle,
with 17—20 segments; aesthetascs on most
segments; accessory flagellum with 4 seg-
ments, terminal segment vestigial.
Antenna 2 (Fig. 2b) about 85% length of
antenna 1. Peduncular segment 4 longer
than peduncular segment 5; peduncular seg-
ment 5 with 3—4 ventral and 2—4 medial
spines. Flagellum with 7 segments, Ist seg-
ment twice as long as average length of
segments 2-6.
Upper lip (Fig. 2c) as long as broad, trap-
ezoidal, with few apical setules.
Mandible (Fig. 2h, 1): Palp 3-segmented,
with 3—4 terminal setae; 2nd segment bear-
ing 2 apical and 2-3 subapical setae. Molar
rounded and well developed, with 1 long
lateral seta. Both incisor and lacinia mobilis
on left mandible with 5 irregular, rounded
cusps (Fig. 2h); 3 long and 3 short plumose
spines between lacinia and molar. Right
mandible (Fig. 21): lacinia apically serrated,
VOLUME 113, NUMBER 1
consisting of irregularly pointed denticles;
2 long and 2 short plumose spines between
lacinia and molar.
Lower lip (Fig. 2d) bearing setules on
outer lobes, outer and mandibular lobes
with rounded corners.
Maxilla 1 (Fig. 2e): Palp 2-segmented,
with 5—6 apical setae and few lateral set-
ules. Outer plate with 6 serrate spines and
1 plumose spine; inner plate with 3 plumose
setae.
Maxilla 2 (Fig. 2f): Outer plate apically
with 1—2 comb-like setae, +15 medium-
sized plumose setae and 2 large plumose
spines (slightly subapical); medial margin
with few fine setules; apical margin of inner
plate bearing +18 long comb-like setae and
+5 short naked setae (seta/spine types in
Fig. 2g).
Maxilliped (Fig. 2j, 6c): Segment 1 of
palp with 1—2 medial setae; segment 2 bear-
ing 12-13 medial setae; dactyl long and
slender, bearing a row of marginal setules.
Inner plates apparently fused along medial
margins.
Gnathopod | (Fig. 3a): Posterior margin
of basis with 6—8 long setae (some doubly
inserted); anterior margin bearing 2—4
spines plus 1 short seta. Carpus with 8 setae
on posterior lobe (4—6 comb-like and 2-3
naked). Propodus ovate, almost twice as
long as broad, larger than gnathopod 2 pro-
podus; palm uneven, serrate with minute se-
tules at corner (Fig. 3b); palmar margin
bearing 7—8 normal spines and 15—19 short
bifid spines on lateral margin; medial mar-
gin with 6 short setae, 1 normal angular
spine, and 4—6 oblique subangular spines
(1-2 relatively long). Dactyl about 70%
length of propodus; inner margin with dis-
tinct row of denticles (Fig. 3c).
Gnathopod 2 (Fig. 3d): Posterior margin
of basis bearing 9-10 long setae. Carpus
posteriorly with fine setules and 7 sets of
setae (1—5 setae per set). Propodus ovate,
almost twice as long as broad; palm
oblique, with 5—6 corner spines (Fig. 3e),
11-14 short lateral bifid spines, and 7-8
short medial setae; palmar margin finely
107
serrate at whole margin, with minute setules
at corner. Dactyl about 50% length of pro-
podus; inner margin with distinct row of
denticles.
Pereopods 3 and 4 subequal (Fig. 4a).
Basis without spines, anterodistal margin
even (pereopod 4 basis with O—1 spine plus
1 seta at anterodistal margin). Posterior
margin of carpus bearing 4—5 spines. Pro-
podus with 8—9 spines along posterior mar-
gin and 2 apical spines. Dactyl about 24%
length of propodus (Fig. 4b).
Pereopod 5 (Fig. 4c): Basis with 10-11
spines at posterior margin (distal and prox-
imal group of spines separated by a gap);
anterior margin bearing 13-15 spines; an-
terior lateral surface with 6 short setae; 10—
11 short setae at posterior margin and pos-
terior lateral surface. Ischium with 1 spine
and 3 setae. Anterior margin of carpus with
a row of long, bifurcate setae (Fig. 4f) and
6—12 spines (some doubly inserted). Pro-
podus anteriorly with a row of long, bifur-
cate setae, occurring progressively shorter
distally, with slightly thicker bases; lateral
margin with 17—19 spines (some doubly in-
serted); proximal part with 3—4 spines. Dac-
ty] 14—20% length of propodus.
Pereopod 6 subequal to pereopod 5 but
slightly longer.
Pereopod 7 (Fig. 4d): Basis ovate, bear-
ing 8 spines on anterior margin and 7—9
spines on posterior margin. Ischium with 2
spines. Merus with 3 spines on posterior
margin and 5 spines at anterior margin (1
singly and 2 doubly inserted). Carpus with
13 spines, occurring in 6—7 sets (with 1-3
spines per set) on anterior margin, 4 spines
(2 doubly inserted) plus 4 setae on posterior
margin, and 10—11 terminal spines. Propo-
dus bearing 12 slender spines on anterior
margin and rows of long, bifurcate setae
along posterior and anterior margins (Fig.
4f). Dactyl about 27% length of propodus.
Pereopods 1—7 without any trace of len-
ticular organs; pereopods 5—7 with broad
bases.
Coxal gills ovate, present on pereopods
4-6.
108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fie:
Brazil.
Coxal plates 1 and 2 (Fig. 3f, g) about
equal in size, subrectangular to ovate in
shape, wider than long; plates 3—6 longer
than wide, overlapping; plates 3 and 4
about the same size, plate 5 largest; plate 5
and 6 with distinct anterodistal lobes (Fig.
4c, g); plate 7 (Fig. 4e) subtriangular to ir-
regular in shape, with single long spine at
tapered posterior corner.
Oostegites linear and elongate, on pereo-
pods 2-5.
Pleopods 1—3 morphologically alike (Fig.
5a), decreasing slightly in size posteriorly,
with subequal outer and inner rami. Outer
rami 3-segmented, with 2 terminal plumose
setae per segment; segment 1 bearing 6—10
lateral plumose setae and 5—8 medial plu-
mose setae. Inner rami 1-segmented, with
4—5 medial plumose setae.
Epimeral (pleonal) plates subquadrate,
with small, subacute posterior margins,
bearing 1 setule each (Fig. 4h).
Uropod 1 (Fig. 5b): Rami subequal in
length, slightly shorter than peduncle. Pe-
duncle with 3 spines on dorsolateral, dor-
Spelaeogammarus spinilacertus n. sp., allotype female (8.1 mm) from Jaburu Cave, Estado do Bahia,
somedial, ventrolateral (basofacial), and
apical margin, respectively. Outer ramus
bearing 3—4 dorsolateral and 4 apical
spines. Inner ramus with 5 apical and 4—5
dorsolateral spines, the latter occurring as
3—4 singly and 1—2 doubly inserted.
Uropod 2 (Fig. 5c): Peduncle bearing 1
dorsomedial spine, 2 dorsolateral spines,
and 2 apical spines. Outer ramus slightly
shorter than inner ramus, as long as pedun-
cle, bearing 2—3 spines dorsolaterally and 4
spines apically. Inner ramus with 5 dorso-
lateral and 4—5 apical spines.
Uropod 3 (Fig. 5d) with subequal, lan-
ceolate rami, both 1-segmented. Peduncle
about 46% length of rami, with 2 apical
spines, 1 subapical spine, and 1 small dor-
soproximal spine. Outer ramus bearing 3
apical spines, 6 sets of spines (with 2-3
spines per set) along lateral margin, and
about 20 bifurcate long setae along medial
margin (Fig. 5e). Inner ramus with 3 apical
spines; lateral margin with 5 spines; medial
margin bearing 9 spines (some doubly in-
serted).
VOLUME 113, NUMBER 1
109
Pig...2.
lower lip, e) maxilla 1, f) maxilla 2, g) spine and seta types (on maxilla | and 2, maxilliped, and gnathopod | and
2), from left: serrate spine, comb-like seta, plumose seta, and plumose spine, h) left mandible, 1) right incisor and
lacinia mobilis, j) maxilliped, k) spine types of maxilliped outer plate (far left 2) and inner plate (far right 2).
Telson (Fig. 5f) width about 84% of
length, with shallow excavation (10% of
length); each side bearing 3 setae, 2 apical
and 2 (sometimes 3) subapical spines.
Etymology.—tThe epithet spinilacertus is
a noun in apposition, alluding to the pres-
&
VODA
Spelaeogammarus spinilacertus n. sp., allotype female: a) antenna 1, b) antenna 2, c) upper lip, d)
ence of spines on the anterior margin of the
basis of gnathopod 1. It is formed by com-
bining spini, from Latin meaning thorn or
spine, with lacertus, from Latin meaning
upper arm (~basis).
Remarks.—Two of the three specimens
110 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Spelaeogammarus spinilacertus n. sp., holotype male: a) left gnathopod 1, b) detail of right gnathopod
1, c) detail of right gnathopod | dactyl, d) left gnathopod 2, e) detail of left gnathopod 2; allotype female: f)
coxal plate of gnathopod 1, g) coxal plate of gnathopod 2.
from Jaburu cave showed variation in some
characters, one of which is diagnostic for
the species: instead of 2—4 spines, the basis
of the female gnathopod 1 had 2-3 setae
along the anterodistal margin (compare Fig.
3a with Fig. 7a and c). These setae had the
same length as the spines they replaced and
could be clearly distinguished from corre-
VOLUME 113, NUMBER 1
11]
Fig. 4. Spelaeogammarus spinilacertus n. sp., allotype female: a) left pereopod 4, b) right pereopod 4 dactyl,
c) left pereopod 5, d) right pereopod 7, e) coxal plate of right pereopod 7, f) bifurcate seta (on carpus and
propodus of pereopod 5-7), g) coxal plate of left pereopod 6, h) epimeral plates.
sponding setae in S. santanensis and S. tra-
jJanoae (described below). Furthermore,
both specimens (10 mm male and 7 mm
female) had 3 subapical spines on each lobe
of the telson. The male from the Jaburu
sample also showed morphological vari-
ability in its appendages, e.g., the bases of
both gnathopods and pereopods 3—6 were
relatively narrow and elongate; similarly,
the propods of both gnathopods appeared
relatively longer and larger, with a conspic-
uously sinusoid palmar margin.
Hi PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
f
Fig: 5:
Spelaeogammarus spinilacertus n. sp., holotype male: a) left pleopod 3, b) left uropod 1, c) left
uropod 2, d) left uropod 3, e) bifurcate seta of outer ramus of uropod 3, f) telson.
Spelaeogammarus trajanoae, new species
Fics. 6b;1, 7c)'d, Sa, b
Material examined.—Campo Formoso,
Estado do Bahia, Brazil: holotype female
(10.4 mm) and 5 paratypes (2 males, 3 fe-
males) from Toca do Pitu Cave (=Gruta do
Pitu?), collected by A. Auler & M. Martins,
Sep 1989; 1 male and | female paratype
from Toca do Pitu Cave, collected by E.
Rubbioli, Jan 1992; 3 paratypes (1 female,
2 juveniles) from Buraco do Teodoro Cave,
collected by J. A. Cardoso (no date given);
1 female paratype from Toca do Gongalo
Cave, collected by P. Gernhard, 4 Jul, 1997;
1 fragmented specimen from Convento
Cave, collected by S. Larizotti, 1986 (?).
The holotype is dissected and mounted
on microscope slides in Faure’s medium.
Holotype (MNRJ 13341) and 2 paratypes
from the type locality are deposited in the
Museu Nacional. The allotype and the re-
maining paratypes are retained in the col-
lection of the second author.
Diagnosis.—Spelaeogammarus traja-
noae is morphologically closely allied with
S. spinilacertus but can be distinguished
from that species by 5—9 short setae instead
of spines or spine-length setae on the dis-
toanterior margin of the basis of gnathopod
1 and coxal plate 5 with approximately 18
setae plus | spine on margins instead of 9
setae plus 1 spine. Largest males 10.0 mm,
largest female 10.5 mm. S. trajanoae is
moreover distinguished from S. spinilacer-
tus as indicated in the following descrip-
tion.
Description.—Antenna 1: Peduncular
segment 1 with 5—7 spines; peduncular seg-
ment 2 with 3—4 short spines; peduncular
segment 3 bearing 2 spines.
Lower lip (Fig. 61) bearing setules on
medial margin of outer lobes.
VOLUME 113, NUMBER |
Maxilliped: Segment 2 with +17 setae
along medial margin. Inner and outer plate
shown in Fig. 6b.
Gnathopod 1 (Fig. 7c): Posterior margin
of basis bearing 9-10 singly inserted, long
setae; anterior margin with 5—9 short setae.
No setules on posterior margin of ischium.
Carpus bearing 6—9 setae on pointed pos-
terior lobe (4—6 comb-like and 2—3 naked).
Palm of propodus even; lateral margin with
5—7 normal spines and 16—17 short bifid
spines; medial margin with 6—8 short and 6
long setae.
Gnathopod 2: Posterior margin of basis
with 8—9 long setae. Carpus posteriorly
with 5—6 rows of setae (1—5 setae per row).
Propodus with 6—8 rows of setae (1—3 setae
per row) at proximoposterior margin; pal-
mar margin with 3—4 lateral corner spines,
16-17 short lateral bifid spines, and 8—9
short medial setae.
Pereopods 3 and 4 (Fig. 8a): Basis bear-
ing | spine at posterodistal margin; antero-
distal margin sinusoid, with 1 spine and 1
seta (pereopod 3) or 2 spines and 1 seta
(pereopod 4). Carpus bearing 6—7 spines
posteriorly. Propodus with 13-15 spines
along posterior margin.
Pereopod 5: Coxal plate with 17—18 mar-
ginal setae on anterior lobe. Anterior mar-
gin of basis bearing 10—12 spines; 15 short
setae on both posterior and anterior lateral
surfaces. Dactyl about 22% length of pro-
podus.
Pereopod 6: Ischium with 2 spines and
1-2 setae. Dactyl about 26% of propodus
length.
Pereopod 7: Basis with 11 spines on an-
terior margin and 10 spines on posterior
margin. Merus with 3 sets of doubly in-
serted spines at posterior margin and 3 sets
of spines at anterior margin (with 3—4
spines per set).
Pleopods: Outer ramus with 8—10 lateral
plumose setae and 8—9 medial plumose se-
tae on first segment. Inner ramus bearing 5—
7 medial plumose setae.
Epimeral plates (Fig. 7d) with produced,
bluntly rounded distoposterior corners.
113
Uropod 1: Peduncle with 3—4 spines on
dorsolateral margin, 4—5 spines on dorso-
medial margin. Inner ramus with 4—5 mar-
ginal spines, occurring as 4 dorsomedial
and O—1 dorsolateral spines.
Uropod 2: Peduncle with 1—2 dorsome-
dial spines, 2—3 dorsolateral spines.
Uropod 3: Outer ramus bearing 3-5 api-
cal spines, 4—6 rows of spines (with 1-3
spines per row) along lateral margin. Inner
ramus bearing 10—11 rows of spines (with
5 singly and 5 doubly inserted) at medial
margin.
Telson (Fig. 8b) width about 81% of
length, with u-shaped apical excavation
(19% of length); each side with 3 apical
setae, 2 apical spines, and 3—4 subapical
spines.
Etymology.—The species is named in
honor of Professor Dr. Eleonora Trajano of
the University of Sao Paulo, who has made
important contributions to Brazilian bio-
speleology.
Remarks.—Out of a total of 12 speci-
mens examined, 2 individuals were found
with 2 and 3 subapical spines on each tel-
sonic lobe, respectively. The majority (10
specimens) had 4 subapical spines on one
lobe and 3 subapical spines on the other
lobe.
Spelaeogammarus santanensis, new
species
Figs. 6a, e—g, 7a, b, 8c—e
Material examined.—Padre Cave, San-
tana, Estado do Bahia, Brazil: holotype
male (13.6 mm), 3 male and 3 female para-
types (11.5—13.6 mm), and | juvenile para-
type (10.6 mm), collected by KE Chaimow-
icz, July 1987.
The holotype is dissected and mounted
on microscope slides in Faure’s medium.
Holotype (MNRJ 13342) and 2 paratypes
are deposited in the Museu Nacional. The
remaining paratypes are retained in the col-
lection of the second author.
Diagnosis.—A comparatively large cav-
ernicolous species, easily distinguished
114 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 6. Spelaeogammarus santanensis n. sp., holotype male: a) inner and outer plate of maxilliped, e) detail
of antenna 1, f) left mandible, g) detail of right mandible; S. trajanoae n. sp., paratype female (10.36 mm): b)
inner and outer plate of maxilliped, i) lower lip; S. spinilacertus n. sp., holotype male: c) inner and outer plate
of maxilliped; S. bahiensis paratype (8 mm juvenile): d) inner and outer plate of maxilliped, h) left incisor and
lacinia mobilis.
VOLUME 113, NUMBER 1
115
Fig. 7.
Spelaeogammarus santanensis n. sp., holotype male: a) left gnathopod 1, b) right gnathopod 2; S.
trajanoae n. sp., paratype female (10.36 mm): c) left gnathopod 1, d) epimeral plates.
from other species in the genus by having:
5-segmented accessory flagellum; 20—23
setae on posterior margins of the bases of
gnathopods 1 and 2; propodus of gnathopod
1 proportionally larger. Largest male 13.6
mm, largest female 10.5 mm. S. santanensis
is furthermore distinguished from S. spini-
lacertus according to the following descrip-
tion.
Description.—Antenna | about 40—45%
length of body. Primary flagellum bearing
20-—21 segments, some of which with mul-
116 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 8.
Spelaeogammarus trajanoae n. sp., paratype female (10.36 mm): a) left pereopod 3, b) holotype
female: telson; S. santanensis n. sp., holotype male: c) right pereopod 3, d) coxal plate of pereopod 7, e) telson;
S. bahiensis, paratype (8 mm juvenile): f) coxal plate of pereopod 7, g) telson.
tiple inserted aesthetascs (Fig. 6e). Pedun-
cular segment 1 with 5—6 spines; pedun-
cular segment 3 with 2 ventromedial spines.
Accessory flagellum 5-segmented.
Antenna 2 slightly shorter than antenna
1. Peduncular segment 5 with 5 ventral
spines. Flagellum as long as peduncular
segment 5, with 8—10 segments.
Mandible (Fig. 6f, g): Palp with 3 ter-
minal setae. Palp segment 2 of left mandi-
ble with 3 setae and 1 spine; 3 short and
1(—2?) long plumose spines between lacinia
and molar. Palp segment 2 of right mandi-
ble with 4 setae and 2 spines.
Lower lip with few thin setules on medial
margin of outer and inner lobes; corner of
outer lobes with slightly pointed corners.
Maxilla 1: Palp with 8—9 apical and sub-
apical setae and a few lateral setules. Outer
plate with 7 serrate spines apically.
Maxilla 2: Outer plate apically with 0-1
comb-like setae, +21 plumose setae, and 1—
2 plumose spines (slightly subapical). Inner
plate bearing +22 long comb-like setae and
VOLUME 113, NUMBER 1
several short naked setae (two of which on
medial margin).
Maxilliped: Palp segment 1 with 2 me-
dial and 2 proximomedial setae; segment 2
with +25 medial setae; segment 3 bearing
+17 medial and 3 dorsomedial setae. Dac-
tyl with 8 setae on outer margin. Inner and
outer plate shown in Fig. 6a.
Gnathopod | (Fig. 7a): Posterior margin
of basis bearing 20 long, singly inserted se-
tae; anterior margin with 4 setae. Carpus
with distinctly pointed posterior lobe,
which bears 13—14 plumose setae. Propo-
dus bearing 7—8 setae on medial surface;
palmar margin sinusoid, with 10 short me-
dial setae, 4 long lateral setae, 8—9 normal
lateral spines, 19-20 short bifid lateral
spines, 1 normal angular spine medially, 2—
3 oblique subangular spines (1—2 relatively
long), +13 subangular lateral setae, and 6—
7 subangular medial setae. Inner margin of
dactyl with row of blunt denticles.
Gnathopod 2 (Fig. 7b): Posterior margin
of basis bearing 21—23 long setae. Carpus
posteriorly with 8—9 rows of setae and 5
short setae distolaterally. Propodus twice as
long as broad; palm with 6 lateral spines
and 1 medial spine, 18 short lateral bifid
spines, 1 long lateral seta, and 12 short me-
dial setae; palmar margin with blunt serra-
tion of whole margin, distinct serration and
minute cilia at corner. Dactyl about 53%
length of propodus, inner margin with row
of blunt denticles.
Pereopod 3: Basis (Fig. 8c) with 2 spines
at posteriodistal margin. Carpus bearing 5—
6 spines posteriorly and 2-3 spines plus 1—
2 setae anteriorly. Propodus with 11-12
spines along posterior margin (some doubly
inserted) and 2 spines plus 4 setae apically.
Dactyl about 25% length of propodus.
Pereopod 4 subequal to pereopod 3, ex-
cept for the following differences: coxal
plate with 1 spine and 8 setae along distal
margin, 2 setae at proximal margin, and 3
setae on lateral surface. Basis with 2—4
spines at posterodistal margin.
Pereopod 5: Coxal plate with 12 setae on
anterior lobe and 2 setae on lateral surface.
i
Basis bearing 12 spines on anterior margin;
anterior and posterior lateral surface with 4
short setae respectively. Ischium with 4 se-
tae. Anterior margin of carpus with 5—6 sets
of spines (2-3 singly and 2-3 doubly in-
serted). Lateral margin of propodus with 13
spines (S—6 doubly inserted). Dactyl about
18% length of propodus, bearing | plumose
seta posteriorly.
Pereopod 6 subequal to pereopod 5 ex-
cept for the following characters: coxal
plate with 1 spine and | seta on anterior
lobe. Basis without setae on posterolateral
surface; anterolateral surface with 2—6 se-
tae; posterior margin bearing 14 spines (pe-
reopod 5: 11 spines). Merus posteriorly
with 3 spines and anteriorly with 3—4 spines
plus 3—4 setae (see S. spinilacertus for pe-
reopod 5). Anterior margin of carpus with
8 spines (3 doubly inserted). Dactyl about
20% length of propodus, bearing 1 plumose
seta and 1 spine posteriorly.
Pereopod 7: Coxal plate (Fig. 8d) irreg-
ular in shape. Basis with 10—11 spines on
anterior margin. Ischium bearing 1 spine
and 1—2 setae. Merus with 6 spines (3 dou-
bly inserted) on anterior margin. Carpus
with 3 sets of spines (with 3—4 spines per
set) on anterior margin and 4 sets of spines
(with 1—3 spines per set) on posterior mar-
gin. Anterior margin of propodus addition-
ally with 10 sets of 1—3 slender spines. Dac-
tyl about 21% length of propodus, with |
plumose seta at posterior margin.
Pleopods: Outer rami bearing 12-13 lat-
eral plumose setae and 7—8 medial plumose
setae. Inner rami with 7—8 medial plumose
setae.
Epimeral plates subquadrate, with small,
subacute posterior margins, bearing | setule
each.
Uropod 1: Peduncle with 4 spines on
dorsolateral, 4 on dorsomedial, 3 on ventro-
lateral, and 3 on apical margin, respectively.
Inner ramus with 7 dorsolateral spines (3-—
4 dorsomedial and 1—2 dorsolateral spines).
Uropod 2: Outer ramus bearing 4 dor-
solateral spines (doubly inserted).
Uropod 3: Outer ramus bearing 2 spines
118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
and 1 seta apically, 5 sets of spines (with
1-3 spines per set) along lateral margin;
medial margin with +8 bifurcate setae
proximally and +6 slender setae distally.
Medial margin of inner ramus with 8 sets
of singly and doubly inserted spines.
Telson (Fig. 8e) width about 76% of
length; apex with shallow excavation (5%
of length); each lobe bearing 1 spine plus
1 seta apically and 3 spines plus 1 seta sub-
apically.
Etymology.—The proposed epithet San-
tanensis is a toponym, referring to the cap-
ital city Santana, which is situated near the
type-locality.
Remarks.—In marked contrast to the 3
other species of the genus, the preserved
specimens of S. santanensis appeared whit-
ish and almost transparent. Specimens of S.
spinilacertus and S. trajanoae were yellow-
ish-grey, whereas S. bahiensis showed a
dark, brownish tone. Since these variations
appeared to be interspecific (i.e., consistent
for species from different localities) it is
possible that they were caused by structural
differences of the exoskeletons.
Spelaeogammarus bahiensis da Silva
Brum, 1975
Figs: 6dhy Tiss
Material examined.—Patamute Cave
(type locality), Curaga, Distrito de Mata-
muté, Estado do Bahia, Brazil: 1 male par-
atype, 11.1 mm, and 1| juvenile paratype, 8
mm (Museu Nacional catalogue no. MN
5725), collected by P Magalhaes, 1972-—
1973 @):
Diagnosis.—Spelaeogammarus bahien-
sis 1s morphologically more similar to S.
spinilacertus and S. trajanoae than to S.
santanensis but differs from the former two
species in having 7 serrate spines on the
outer plate of maxilla 1, 20—21 setae on
coxal plate 5, and the presence of dorsal
setules on pereonite 7 (1 setule), pleonites
1—3 (2-10 setules), and uronites 1—2 (1-7
setules). In addition, it is distinguished from
S. spinilacertus by short setae instead of
spines on the anterior margin of the basis
of gnathopod 1. Male specimen 11.1 mm in
length. Corresponding to the original de-
scription by da Silva Brum (1975) with the
additions and modifications given below.
Redescription.—Antenna 1: Peduncular
segment 1—3 gradually decreasing in length;
peduncular segment 1 with 6 ventral spines.
Aesthetascs on most segments of primary
flagellum (as long as segments); accessory
flagellum with 4 segments, terminal seg-
ment vestigial.
Antenna 2: Peduncular segment 3 with
3—4 spines. Flagellum with 7 segments.
Mandible: Left lacinia mobilis distinctly
toothed. Long and short plumose spines be-
tween lacinia and molar more dissimilar
than in other species (Fig. 6h).
Maxilla 1: Palp without lateral setules.
Outer lobe with 7 apical spines (3 different
types) and distinct row of setules at medial
margin, subapically.
Maxilliped: Dactyl bearing 3—4 relatively
long setae along inner margin. Inner plate
bearing apically 2 bifid (y-shaped) spines;
outer plate with 3 blade-like spines sub-
apically and 2 strong setae apically (Fig.
6d).
Gnathopod 1: Posterior margin of basis
with 7—9 long setae (all singly inserted); an-
terior margin bearing 3—5 short setae. Car-
pus with rectangular posterior lobe, bearing
3-5 long, sickle-shaped, naked setae, 3—6
comb-like setae, and 3—4 long, naked setae.
Palmar margin of propodus bearing 5-9
normal spines and 13—18 short bifid spines
on the lateral margin; medial margin with
8—10 short setae, 1 normal angular spine,
and 3—5 corner spines (1—2 relatively long).
Dactyl with 9-10 denticles, each with 1
short seta.
Gnathopod 2 basically like that of S. spi-
nilacertus. Palp with 2 normal spines and 4
spines at corner. Dactyl with 6—7 denticles.
Pereopods 3 and 4 subequal. Pereopod 3
basis with | distoanterior spine, and 1 dis-
toposterior seta. Pereopod 4 basis with 1—2
distoanterior spines and | seta plus 1 spine
VOLUME 113, NUMBER |
at distoposterior margin; dactyl with | spine
similar to S. santanensis.
Pereopods 5—7 missing in specimens ex-
amined.
Pleopods basically like those of S. spi-
nilacertus. Segment 1 bearing 7-8 lateral
plumose setae and 5—6 medial plumose se-
tae. Inner ramus with 7 plumose setae on
medial margin.
Coxal plates 3—4 with 9-10 setae and 1
spine; plates 5—6 identical, bearing 20—21
setae and 1 spine, respectively; plate 7 (Fig.
8f) irregular, slightly lobed.
Uropod 1: Peduncle with 4 spines on
dorsolateral, dorsomedial, and ventrolateral
margins, respectively, and 2 on apical mar-
gin. Outer ramus with up to 5 apical spines.
All spines on inner ramus singly inserted.
Uropod 2: Peduncle bearing 1—2 dorso-
medial, 1—2 dorsolateral, and 2 apical
spines.
Uropod 3 missing in specimens exam-
ined.
Telson (Fig. 8g) width about 80% of
length; each lobe bearing 2 setae, 2 apical
and 3—4 subapical spines.
Discussion
The four species described above are re-
corded from eight caves in a series of dis-
continuous karst areas that extend over a
linear distance of ca. 1200 km from north
to south in eastern Brazil (Fig. 9). The rec-
ords for each species per karst area are as
follows: S. santanensis—1 cave, S. bahien-
sis—1 cave, S. spinilacertus—2 caves, and
S. trajanoae—4 caves. Each species is ob-
viously restricted to one or more caves in a
separate karst area. These areas are disjunct
and apparently physically isolated from
each other. The greatest distance between
caves with two species is roughly 650 km
(S. bahiensis and S. santanensis), whereas
the shortest distance is only about 135 km
(S. bahiensis and S. spinilacertus). Caves
situated within a single, continuous karst
area that are inhabited by the same species
are never more than ca. 100 km apart.
119
Both the elongate coxal plates and ae-
quiramus pleopods found in the genus Spe-
laeogammarus are characters usually con-
sidered plesiomorphic for the family Bogi-
diellidae (Stock 1981, Barnard & Barnard
1983). Apart from Spelaeogammarus, coxal
plates that are longer than wide are known
only for the genus Artesia Holsinger, 1980
(in Holsinger & Longley 1980), and this ge-
nus, like Spelaeogammarus, also has pleo-
pods with aequiramus inner rami. However,
they are 5-segmented in Artesia and only 1-
segmented in Spelaeogammarus. Artesia
can also be distinguished from Spelaeo-
gammarus by 1-segmented accessory fla-
gellum, fewer flagellar segments in both an-
tennae, l-segmented palp of maxilla 2, 6-
segmented pleopodal exopodite, unlobed
coxal plates 5 and 6, and the telson, which
is deeply cleft and bears 4—6 apical spines
on each lobe. Equally long rami are also
found in Aequigidiella Botosaneanu &
Stock, 1989, Kerguelenicola Ruffo, 1974,
and Parabogidiella Holsinger, 1980 (in
Holsinger & Longley 1980). However, in
these three genera the coxal plates are typ-
ically wider than long. Moreover, Aequigi-
diella differs from Spelaeogammarus by
sexually dimorphic inner rami and spines of
the second uropods of the male and a telson
that is much longer than wide. Although to
date only one specimen of the genus Ker-
guelenicola is known, there are several
characters that distinguish it from Spelaeo-
gammarus: 1-segmented accessory flagel-
lum, distinct shape of and lack of armature
on the telson, large mandibular molar, and
reduced number of spines and setae on the
outer and inner lobes of maxilla 1. Para-
bogidiella differs from Spelaeogammarus
by l-segmented accessory flagellum, 5-seg-
mented flagellum of antenna 2, 1-segment-
ed palp of maxilla 2, characteristically elon-
gated pereopod 7, 5 pairs of coxal gills, and
armature of the telson.
The most closely related bogidiellid tax-
on to Spelaeogammarus described to date
may be Bogidiella gammariformis Sket
(1985) from a cave in Equador. This species
120 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
VENEZUELA
bd San —_
>
COLOMBIA Bi
ia .
Pacific
Ocean
Fig. 9:
BRAZIL
’ Salvador
2 mi
Rio De Janeiro
Sao Paulo
Atlantic Ocean
Distribution of species of Spelaeogammarus in eastern Brazil: 1) S. bahiensis (1 cave); 2) S. trajanoae
(4 caves); 3) S. spinilacertus (2 caves); 4) S. santanensis (1 cave). Shading indicates principal cave and/or karst
areas. Map adapted from Trajano & Sanchez (1994).
features some interesting characters that
might be interpreted as intermediate states
between the relatively primitive Spelaeo-
gammarus and the more derived Bogidiella
s. str. For example, B. gammariformis has
enlarged, bilobed coxal plates 5 and 6,
which are longer than wide, therefore show-
ing a strong resemblance to the coxae of
Spelaeogammarus. The inner rami of the
pleopods of B. gammariformis are also 1-
segmented, but show the same reduction as
in most other species of Bogidiella s. str.
(i.e., shorter than segment 1 of the outer
ramus).
The concentration of the four morpho-
logically closely similar species of Spelaeo-
a
Fig. 10. Distribution of bogidiellid genera in continental South America: 1) Bogidiella cooki Grosso &
Ringuelet, 1979; 2) B. gammariformis Sket, 1985; 3) B. neotropica Ruffo, 1952; 4) B. (Dycticogidiella) ringueleti
Grosso & Fernandez, 1988; 5) B. (Dyct.) talampayensis Grosso & Claps, 1985; 6) B. (Mesochthongidiella)
VOLUME 113, NUMBER 1 121
sein Pa ss
pie
10S
- “St
20S
Pacific Ocean
30S
ARGENTINA Atlantic Ocean
40S
50S
80W 7OW 60W SOW 40W
tucumanensis Grosso & Fernandez, 1985; 7) B. (Stygogidiella) hormocollensis Grosso & Fernandez, 1988; 8)
B. (Styg.) lavillai Grosso & Claps, 1984; 9) Eobogidiella purmamarcensis Karaman, 1982; 10) Marigidiella
brasiliensis Stock, 1981; 11) Megagidiella azul Koenemann & Holsinger, 1999; 12) Patagongidiella danieli
Grosso & Fernandez, 1993 and P. mauryi Grosso & Fernandez, 1993 (sympatric species); 13) Pseudingolfiella
chilensis Noodt, 1965; 14) Spelaeogammarus bahiensis da Silva Brum, 1975, S. santanensis n. sp., S. spinila-
certus n. sp., and S. trajanoae n. sp.
[22
22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
gammarus in a series of disjunct caves is
unique for South America (see Fig. 10).
Particularly interesting are the relatively
subtle morphological differences between
the four species that, in turn, appear to be
correlated with the interspecific spatial dis-
tance as well as the abundance of species
per area: S. spinilacertus and S. trajanoae
occur in the central part of the range of Spe-
laeogammarus and show the highest mor-
phological resemblance. In contrast are the
more obvious differences between the rel-
atively large S. santanensis and the smaller
S. bahiensis, which occur on opposite ends
of the generic range.
In South America there is a second con-
centration of species in northern Argentina,
which occur exclusively in hyporheic (7?)
habitats along the Rio Grande (see Fig. 10).
This cluster is also strictly endemic but it
has a higher generic diversity, with six spe-
cies in two genera and three subgenera,
possibly reflecting the time of divergence
from a very old freshwater precursor and
the subsequent radiation into a region of
isolated inland habitats. However, the dis-
tribution pattern of Spelaeogammarus
seems to indicate quite a different historical
scenario, inasmuch as the species of this ge-
nus show an exceptionally close morpho-
logical relationship with each other when
compared with all other South American
bogidiellids. If we assume, for the sake of
argument, the same evolutionary rate for all
South American bogidiellids, the cluster of
species belonging to Spelaeogammarus ap-
pears to have originated from a common
ancestor far more recently as opposed to
other bogidiellids in South American fresh-
water habitats.
The morphological appearance of the
four species as well as their distribution
Over a relatively wide range of disjunct
karst “‘islands’”’ characterize Spelaeogam-
marus as a distinct genus within the family
Bogidiellidae. The apparent isolation of
these species may well reflect a sequence of
allopatric (geographic) speciation events
over a relatively short period of time.
Acknowledgments
We are grateful to Professor Eleonora
Trajano for providing us with some of the
specimens examined in this study and for
helpful comments on Brazilian caves and to
Dr. Paolo S. Young, curator of the Museum
Nacional, Rio de Janeiro, for making the
paratypes of Spelaeogammarus bahiensis
available to us. This study was supported
by a PEET grant from the National Science
Foundation to JRH (DEB-9521752). We
thank the Graphic Office at Old Dominion
University for assistance with preparation
of the distribution maps.
Literature Cited
Barnard, J. L., & C. M. Barnard. 1983. Freshwater
Amphipoda of the world.—I. Evolutionary pat-
terns. Hayfield Associates: Mt. Vernon (Virgin-
ia).
Botosaneanu, L., & J. H. Stock. 1989. A remarkable
genus of cavernicolous Bogidiellidae (Crusta-
cea, Amphipoda) from Thailand.—Studies in
honour of Dr. Pieter Wagenaar Hummelinck.
Foundation for Scientific Research in Surinam
and the Netherlands Antilles, Amsterdam, no.
123;
da Silva Brum, I. N. 1975. Spelaeogammarus bahien-
sis g.n. sp.n. de Anfipodo Cavernicola do Brasil.
(Amphipoda-Bogidiellidae).—Atas da Socieda-
de de Biologia di Rio de Janeiro 17:125—128.
Grosso, L. E., & G. L. Claps. 1984. Tercer Bogidiel-
lidae (Crustacea Amphipoda) de la cuenca del
Rio Grande (Jujuy, Argentina).—Neotropica
30:223-231.
ee ss . 1985. Distribucién geogrdafica de
la familia Bogidiellidae (Crustacea, Amphipo-
da) en la Republica Argentina, con la descrip-
cidn de un nuevo subgénero y una nueva es-
pecie.—Physis (Buenos Aires), Secc. B 43:49—
523
, & H. R. Fernandez. 1985. Una nueva Bogi-
diella (Amphipoda Bogidiellidae) hiporreica de
la provincia de Tucuman (Argentina).—Neotro-
pica 31:201—209.
pide . 1988. Un caso de simpatria de tres
especies del género Bogidiella (Crustacea, Am-
phipoda) en el noroeste Argentino, con la de-
scription de dos nuevoas especies.—Stygologia
4:64-78.
Pee . 1993. Nuevo género cavernicola
austral de Bogidiellidae; Patagongidiella n.
gen. del noroeste Patagonico (Neuquén, Argen-
VOLUME 113, NUMBER 1
tina).—Bolletino del Museo Civico di Storia
Naturale Verona 17:357-372.
, & R. A. Ringuelet. 1979. Fauna subterranea
de las aguas dulces de la Republica Argentina.
I. Dos nuevas especies de Amphipodos del gé-
nero Bogidiella.—Limnobios 1:381—394.
Holsinger, J. R., & G. Longley. 1980. The subterranean
amphipod crustacean fauna of an artesian well
in Texas.—Smithonian Contributions to Zoolo-
gy 308:1-62.
Karaman, G. S. 1982. Critical remarks to the recent
revisions of bogidiella-group of genera with
study of some taxa (fam. Gammaridae). Contri-
bution to the knowledge of Amphipoda 126.—
Poljoprivreda I Sumarstvo Titograd 28:31—57.
Koenemann, S., & Holsinger, J. R. 1999. Megagidiella
azul n. gen., n. sp., a cavernicolous amphipod
crustacean of the family Bogidiellidae from
Brazil, with remarks on its biogeographic and
phylogenetic relationships.—Proceedings of the
Biological Society of Washington 112:572—580.
123
Noodt, W. 1965. Interstitielle Amphipoden der kon-
vergenten Gattungen I/ngolfiella Hansen und
Pseudingolfiella n. gen. aus Suedamerika.—
Crustaceana 9:17—30.
Ruffo, S. 1952. Bogidiella neotropica n. sp., nuovo
Anfipodo dell’ Amazonia.—Rivista Svizzera di
Idrologia 14/1:129-—134.
. 1974. Nuovi Anfipodi interstiziali delle coste
del Sud Africa.—Atti dell’ Istituto Veneto di
Scienze, Lettere ed Arti 132:399-—419.
Sket, B. 1985. Bogidiella (s. 1.) gammariformis sp. n.
(Amphipoda) from Equador.—BioloSk, Vestnik
33:81-88.
Stock, J. H. 1981. The taxonomy and zoogeography
of the family of Bogidiellidae (Crustacea, Am-
phipoda), with emphasis on the West Indian
taxa.—Bijdragen tot de Dierkunde 51:345-—374.
Trajano, E., & L. E. Sanchez, 1994. Bresil. Pp. 527-
540 in C. Juberthie, & V. Recu eds., Encyclo-
paedia Biospeologica, Tome I.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):124—128. 2000.
Eudendrium bathyalis, a new species of hydroid
(Hydrozoa: Anthomedusae: Eudendriidae) from Bermuda
Antonio C. Marques and Dale R. Calder
(ACM) Departamento de Biologia - Faculdade de Filosofia, Ciéncias e Letras de Ribeirao Preto -
Universidade de Sao Paulo. Av. Bandeirantes 3900, 14040-9001, Ribeirao Preto, SP, Brazil.
E-mail: marques @ ffclrp.usp.br;
(DRC) Centre for Biodiversity and Conservation Biology, Invertebrate Zoology, Royal Ontario
Museum, 100 Queens Park, Toronto, Ontario, MS5S 2C6, Canada. E-mail: dalec@rom.on.ca
Abstract.—Eudendrium bathyalis, a new species, is described from relatively
deep-water (283 m) on the offshore slope of the oceanic island of Bermuda.
Its distinctive characters are the small size of the colony, the cnidome, com-
prising two sizes of microbasic euryteles and the unreduced female blastostyle,
bearing eggs supported by long peduncles.
The shallow-water hydroids of Bermuda
were recently revised in a series of studies
by Calder (1988, 1991, 1997). These: re-
ports complemented several earlier studies
on the fauna of the region (viz. Allman
1888, Congdon 1907, Ritchie 1909, Bennitt
1922, among others).
Combining records from these studies
with collections of hydroids from deeper
waters (Calder 1996, 1998), a total of 110
species has been reported from this small
oceanic island. The relatively large number
of species from a small geographic area
likely reflects the extensive sampling un-
dertaken in the area, from the intertidal
zone to abyssal bottoms, including bays,
grassbeds, caves, reefs, ponds, mangroves,
and pelagic seaweeds.
Within the Eudendriidae, four species be-
longing to the two known genera of the
family (Myrionema amboinense Pictet,
1893, Eudendrium bermudense Calder,
1988, Eudendrium capillare Alder, 1856,
and Eudendrium carneum Clarke, 1882) are
known from Bermuda.
These four species appear to be wide-
spread in the warm western Atlantic. An-
other species recorded from the Caribbean
region is Eudendrium ramosum (Linnaeus
1758) (Wedler 1975, for Colombia; Wedler
& Larson 1986, for Puerto Rico); this spe-
cies is also present in the fauna of Bermuda
(pers. obs.). Other species of Eudendrium
have also been described or reported for the
region (Allman 1877, Fraser 1944). These
species are poorly known taxonomically,
being characterized on gross morphology
alone; in some cases, hydranths were lack-
ing, and no information exists on their com-
plement of nematocysts.
The purpose of this report is to describe
a new species of Eudendrium, collected by
submersible from bathyal waters on the
slope of the Bermuda Pedestal.
Methods
Material was collected by submersible
(SDL-1) on 3 Mar 1997, depth 283 m, from
the Bermuda Pedestal. The study area is
discussed more fully elsewhere (Calder
1998). The holotype was examined, mea-
sured, and drawn under microscope and ste-
reomicroscope, both with camara lucida.
The cnidome terminology follows Weill
(1934) and Mariscal (1974), and measures
of nematocysts were made on non-dis-
charged capsules. The L/W ratio (Kubota
1976) and S/C ratio (Watson 1987) are also
provided. Other study methods for Euden-
VOLUME 113, NUMBER 1
driidae are from Marques (1995) and
Marques & Migotto (1998).
Systematics
Genus Eudendrium Ehrenberg, 1834
Eudendrium bathyalis, new species
Fig. 1
Examined material.—Holotype, one fe-
male colony, Bermuda Pedestal 32°16.6'N
ma 44.5 W, 283 m,; 3 Mar 1997, on a
sponge, ROMIZ B3034.
Description.—Colonies dioecious, frag-
ile, up to 18 mm in height; main stems
slightly fascicled basally or up to the half
of the colony, sometimes formed only by a
couple of tubes, fascicled region up to 0.18
mm in diameter. Hydrocauli arising from
stolonal hydrorhiza growing over a sponge;
branches few, irregular, occurring over en-
tire hydrocaulus, branches up to third order,
in radiate planes; pedicels arising from
main stem or branches of first and second
order. Perisarc of main stem weakly devel-
oped, single tubes 0.08—0.10 mm in diam-
eter, unfascicled region with scarce annu-
lations, in sets of 2—3 rings. Branches with
2-5 rings at origin, 0.06—0.08 mm in di-
ameter. Pedicels obscurely annulated at or-
igin, with 2—4 rings, very delicate, 0.05—
0.06 mm in diameter.
Hydranths 0.10—0.25 mm in height,
0.10—0.29 mm in diameter (measured in the
body region just below the tentacles), with
a distinct deep groove in the aboral region;
hypostome large; tentacles 18—24 in num-
ber, occurring in a whorl below hypostome.
Gonophores styloids, arising from body
of hydranth. Immature styloids placed in a
circle around body of hydranth. Female
blastostyles styloids without a characteristic
Spadix over a single egg. Tentacles and hy-
postome not reduced during ontogeny of fe-
male gonophores. Eggs almost circular, ma-
turity undeterminable, encapsulated by a
thin gelatinous layer, linked by long pedun-
cles to body of hydranth, distal part of pe-
duncle broadened for egg support. Eggs 3-—
125
5 in number, 0.14—0.15 mm in diameter.
Male gonophores not observed.
Nematocysts of one category, heterotri-
chous microbasic euryteles in two size clas-
ses.
Small microbasic euryteles (not seen dis-
charged), 6.0-6.3 by 2.8-3.2 wm, L/W =
1:2.0—2.1, oval, abundant; distributed over
hydranth body, hypostome, peduncle coe-
nosarc of female gonophore, and tentacles.
Large microbasic euryteles (seen dis-
charged), 22.1—23.4 by 8.2—-8.9 um, L/W =
1:2.6—2.7, bean-shaped; shaft crossing
about from 0.5 to 0.75 of the whole length
of undischarged capsule, discharged shaft
heavily armed, ca. 19.8 ym in length, pro-
portion S/C = 1.4; nematocysts distributed
over hydranth body (sometimes common),
egg peduncle (rare), and coenosarc (not
seen on hypostome).
Etymology.—bathyalis, adj. from Greek
bathys = deep, in allusion to the bathyal
depth from which material of this species
was collected.
Remarks.—The distinctive character of
this species is the pedunculated gonophore,
which we presumed would be a female one.
A similar female gonophore has only been
observed in Eudendrium vervoorti Marques
& Migotto, 1998, a recently described spe-
cies based on scarce material, without hy-
drorhiza and covered by debris, from the
coast of the Netherlands. In E. vervoorti,
the gonophores apparently arose from the
hydranth body, without a conspicuous spa-
dix; some other gonophores (presumably
older ones) are linked to the body of hy-
dranth or to the pedicel by long stalks
(Marques & Migotto 1998). Another simi-
lar feature between both species is the ex-
tremely delicate branches (widths barely
greater than 0.1 mm), but this character is
also shared by several other species of the
genus [e.g., Eudendrium album Nutting,
1896, Eudendrium fragile Motz-Kossows-
ka, 1905, Eudendrium generale von Len-
denfeld, 1885, Eudendrium tottoni Ste-
chow, 1932 (=Eudendrium antarcticum
Totton, 1930, see Stechow 1932) and Eu-
126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
A
Fig. 1.
y
C D E
Eudendrium bathyalis, new species; all from holotype ROMIZ B3034. A, hydranth with the distri-
bution of the large microbasic euryteles represented over the body; B, female blastostyle; C, capsule of the large
microbasic eurytele; D, discharged large microbasic eurytele; E, capsule of the small microbasic euryteles. Scale
bar, A-B = 0.1 mm; C—E = 10 pm.
dendrium motzkossowskae Picard, 1951],
but none of those has a female gonophore
as in E. vervoorti and E. bathyalis.
The cnidome is the distinctive character
between E. vervoorti and E. bathyalis. The
Netherlands species has only small micro-
basic euryteles and the species from Ber-
muda has both small and large nematocysts.
The material from Bermuda is slightly fas-
cicled, but as the Netherlands material is
represented only by a small fragment, even
without hydrorhiza (Marques & Migotto
1998), this difference could be only an ar-
tifact.
Key to the species of Eudendriidae
recorded from Bermuda
la. More than 40 tentacles arranged in two
whorls, presence of abundant zooxan-
thellac-4c nn. Ss Myrionema amboinense
lb. Usually fewer than 35 tentacles ar-
ranged in a unique whorl, zooxanthel-
las absent th) 3-4": Se eee 2
VOLUME 113, NUMBER 1
2a. Nematocysts of only one size class, mi-
crobasic eurytele type
“65 OT SS Cee ee Eudendrium capillare
2b. Nematocysts in two size classes .... 3
3a. Cnidome including small and large mi-
Sea CUR YUCICS 5 ox. ys. spite w wy etailers +
3b. Cnidome including small microbasic
euryteles and large nematocysts differ-
ent from microbasic euryteles ...... 5
4a. Female gonophore encircled by un-
mumimed Spagix'. 27 1. 8S 70S
4b. Female gonophore supported by long
peduncles, no characteristic spadix .
ae Eudendrium bathyalis, new species
5a. Large nematocysts heterotrichous ani-
sorhiza, female immature spadix bifid
Rete. kak h ans Eudendrium carneum
5b. Large nematocysts macrobasic euryte-
les, female immature spadix un-
ie i eee
ofl Eudendrium bermudense
Acknowledgments
The authors wish to thank to Prof. Wim
Vervoort (Nationaal Natuurhistorisch Mu-
seum, Leiden, The Netherlands), in which
laboratory part of the study was done, to
Dr. Alvaro E. Migotto (University of Sao
Paulo, Sao Sebastiao, Brazil) and Dr. Jean-
ette E. Watson (Museum of Victoria, Aus-
tralia) for suggestions concerning the man-
uscript, and to Charles M.D. Santos for his
help with some drawings. ACM has finan-
cial support from the Fundagao de Amparo
a Pesquisa do Estado de Sao Paulo (FA-
PESP 96/10544-0, 97/04572-4, and 98/
14822-0). DRC acknowledges the Canadian
Navy for the opportunity to dive in the
SDL-1 submersible, used in collection of
this hydroid. Thanks are also due to the
Natural Sciences and Engineering Research
Council of Canada for financial support.
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Alder, J. 1856. A notice of some new genera and spe-
cies of British hydroid zoophytes.—Annals and
Magazine of Natural History, 2nd series 18:
353-362.
127
Allman, G. J. 1877. Report on the Hydroida collected
during the exploration of the Gulf Stream by
L.F de Pourtalés, assistant United States Coast
survey.—Memoirs of the Museum of Compar-
ative ZoGlogy at Harvard College 5(2):1—66.
. 1888. Report on the Hydroida dredged by
H.M.S. Challenger during the years 1873-76.
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128 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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opment). II. Le valeur taxonomique du cni-
dome.—Travaux de la Station Zoologique de
Wimereux 11:1—351.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):129-144. 2000.
Cnidae of two species of Discosomatidae
(Cnidaria: Anthozoa: Corallimorpharia) from Brazil
Suzana Machado Pinto and Maria Julia da Costa Belém
(SMP) Depto. de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro/
UFRJ, Quinta da Boa Vista, 20940-040, Rio de Janeiro, RJ.; Current address:
Depto. de Zoologia, Instituto de Biociéncias, Universidade de Sao Paulo (USP), Caixa Postal
11461, 05422-970. Sao Paulo, SP, Brasil. e-mail: suzanamp @ib.usp.br;
(MJCB) Caixa Postal 24.030, 20.522-970. Rio de Janeiro, RJ, Brasil.
e-mail: belcosta@ mandic.com.br
Abstract.—The cnidae of Discosoma carlgreni (Watzl, 1922) and Discosoma
sanctithomae (Duchassaing & Michelotti, 1860) are described and illustrated.
Five types (sensu Schmidt 1969, 1974) were observed: spirocysts, b-rhabdoids,
p-rhabdoids D, holotrichs I and holotrichs II. An unusual variety of holotrich
occurred in the tentacles and column that had not been reported previously.
These species are distinguished based on the distribution and size of the types
of cnidae. The importance of qualitative studies of corallimorpharians cnidae
is discussed.
This paper investigates the diversity of
cnidae that occur in the corallimorpharians
Discosoma carlgreni (Watzl, 1922) and D.
sanctithomae (Duchassaing & Michelotti,
1860) from Brazil, as well as their taxo-
nomic value.
The diagnostic value of cnidae in differ-
ent structures of cnidarians has been ex-
amined by several authors (e.g., Weill 1934,
Russell 1938, Carlgren 1949, Cutress 1955,
Schmidt 1972, 1974, Mariscal 1974, den
Hartog 1980, Fautin 1986, England 1991,
Williams 1996, Pires 1997, Ostman & Hyd-
man 1997).
Corallimorpharia is a relatively small or-
der of skeletonless Anthozoa, morphologi-
cally intermediate between sea anemones
and corals. Many authors consider them
most closely related to corals (e.g., Duerden
1904, Stephenson 1921, Schmidt 1974, den
Hartog 1980, Fautin & Lowenstein 1992,
Chen et al. 1996, Pinto & Belém 1997, Pi-
res & Castro 1997). Contributions to the
systematics of the Discosomatidae, based
on morphological and anatomical charac-
ters, have been made by several authors
(e.g., Duchassaing & Michelotti 1864,
Duerden 1900, Stephenson 1921, Carlgren
1940, 1949, Corréa 1964, Schmidt 1972,
1974, den Hartog 1980, Schlenz & Belém
1982). The revision of the Caribbean shal-
low-water Corallimorpharia, as set forth by
den Hartog (1980), provided a comprehen-
sive review of the morphology, anatomy,
histology and classification of this group.
den Hartog (1980) rearranged the family
Discosomatidae Duchassaing & Michelotti,
1864 on the basis of an array of material
from the Caribbean. He united the five gen-
era of Actinodiscidae (Carlgren, 1949) in
the genus Discosoma Riippell & Leuckart,
1828. den Hartog (1980) concluded that the
order Corallimorpharia does not fundamen-
tally differ from corals and should be in-
cluded as a separate sub-order in the Scler-
actinia.
The cnidom of the Discosomatidae was
previously studied by Watzl (1922), Carl-
gren (1927, 1949), Corréa (1964), den Har-
tog (1980) and Schlenz & Belém (1982),
but these studies were not sufficiently de-
tailed to provide a good understanding of
the cnidae in this group.
130 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Materials and Methods
Specimens of Discosoma carlgreni (Fig.
1A) and D. sanctithomae (Fig. 1B) were
collected from 1990 to 1993 and deposited
in the Cnidaria collection of the Museu Na-
cional/Universidade Federal do Rio de Ja-
neiro, Rio de Janeiro (MNRJ). For this
study, 57 specimens of D. carlgreni and 42
specimens of D. sanctithomae were exam-
ined. Both species were also acquired by
exchange with Mexico and Cuba in order
to carry out a comparative study. Speci-
mens of D. carlgreni were found at the
South Coast of the state of Espirito Santo
and Bahia. So far, D. sanctithomae has been
found only in Abrolhos Archipelago, be-
tween the channel of Redonda and Siriba
Islands (see Appendix for details).
Material was collected by snorkel and
SCUBA diving at depths between | m and
16 m, using a hammer and chisel. Living
animals were placed in plastic sacks with
water from their site of collection and trans-
ported on ice. Specimens were anesthetized
by slowly adding 1:1 solution of 8% MgCl,
and sea water. After anesthetization, they
were fixed in 4% formalin. Cnidae were
studied in squash preparations of living and
preserved specimens. A fragment of tissue
was squashed and spread on a slide. Five
structures were examined: discal tentacle,
marginal tentacle, column, stomodaeum
and mesenterial filament. Measurements
were made only on undischarged capsules.
At least 40 length/width measurements
were made of each nematocyst type in each
structure examined. Measurements were
made with light microscopy at magnifica-
tions up to 1250 with differential inter-
ference contrast optics (Nomarski), using
an eyepiece micrometer. Drawings were
made with a camera lucida. Cnidae were
classified according to the nomenclatures of
Weill (1934, later modified by Carlgren
1940), Schmidt (1969, 1972, 1974) and den
Hartog (1980).
Results
Five types of cnidae were observed: spi-
rocysts, b-rhabdoids, p-rhabdoids D, holo-
trich I and holotrich II. The cnidae were
classified as follows.
Spirocysts (Figs. 3A, 6E).—Elongate and
thin-walled capsule with a long tube coiled
in numerous spirals, tubule without spines.
Remarks.—This variety was sparse but
typical of the marginal tentacles of both
species. The capsules reached up to 18 wm
in length and 3.1 wm in width. Eight un-
discharged capsules in each species were
observed. They were not included in Table
1 due to their sparseness.
B-rhabdoids.—Varied in shape and size,
capsules generally oval in aspect. The
thread of the b-rhabdoids is always armed
with spines, not having a clear difference
between the width of the proximal part of
the tubule and that of the distal portion. We
observed two morphological varieties of b-
rhabdoids in undischarged capsules:
B-rhabdoids' (Figs. 2B, 3C, 4B, 6K 7B,
Table 1).—Capsule oval to cylindrical in
shape, of refractive contrast. In the undis-
charged state, the basal portion of the tu-
bule is shorter and thinner than that of b-
rhadoids’, up to 0.33 of the length of the
capsule. Sometimes the basal part appears
slightly curved, the tubule being arranged
in few irregular coils.
Remarks.—The undischarged capsules
from the column may be occasionally con-
fused with holotrich II, due to little refrac-
tive contrast.
B-rhabdoids* (Figs. 2A, 3B, 4A, Table
1).—Capsule transparent, elongate, very
small and of refractive contrast. Its basal
portion is short, refractive and clear.
P-rhabdoids D.—Characterized by a no-
tably wide tubule ending with a funnel-
shaped “‘V”’ with two distinct parts: large
basal shaft and tapered distal tubule. The
thread is long, bearing spines. We observed
two morphological varieties of p-rhabdoids
D in undischarged capsules:
P-rhabdoids D' (Figs. 2C, D, 3D, 4C, D,
VOLUME 113, NUMBER 1 131]
Fig. 1. A, Discosoma carlgreni. Specimens collected among Zoanthus sp. at Santa Cruz, Aracruz, ES. Scale:
5 cm. B, D. sanctithomae. Aggregation from study site at Abrolhos Archipelago, between the channel of Redonda
and Siriba Islands. Scale: 2 cm.
132 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
G, 5B, C, 6A, B, G, H; 7A, H; 8A, B; Table
1).—Capsule cylindrical, slightly curved, of
refractive contrast. The shaft has obvious
turns of spines that are up to 0.5 of the cap-
sule length.
Remarks.—This variety was rather com-
mon and also occurred in all structures ex-
amined in both species. It also occurred in
two size classes in the discal tentacles, col-
umn and mesenterial filaments. In the fila-
ments of D. sanctithomae some transparent
thin-walled capsules, with rich contrast in
the shaft, were observed (Fig. 8B).
P-rhabdoids D? (Fig. 7D, Table 1).—Rel-
atively large elongate capsule, cylindrical,
slightly curved and of refractive contrast.
Shaft with clear turns of spines can reach
up to 0.5 length of the undischarged cap-
sule. A long tubule is irregularly arranged
inside the capsule.
Holotrichs—We observed two morpho-
logical varieties of holotrich in both spe-
cies:
Holotrichs I (Figs. 2E, G, 3E G, 4EF H,
I, 5A,.Ds 6G. D, 1 Ko IE, BGI se.
Table. 1).—Capsules of various sizes, filled
with a tubule having a long figure eight
form, of rich contrast in several turns (Figs.
2G, 6K). In the discharged state, the tubule
contains conspicuous spines of equal size
distributed along almost its entire length,
except for a short, naked basal portion. The
distal end of the tubule presents an abrupt
tapered distal tip, completely devoid of
spines and measuring up to 30 pm (Figs.
2G, 6D).
Remarks.—The distal end of the tubule
of these nematocysts was previously de-
scribed by den Hartog (1980) as a terminal
tubule. The terminal tubule can be seen
only when the tubule is totally discharged.
Holotrichs I, which are common, occurred
in two size classes throughout all structures
examined, except for the presence of only
small capsules in the column of D. carl-
greni (Fig. 4E Table 1). This type ranges in
size from the small (e.g., in marginal ten-
tacles—25-—34 by 9.4—18 ym) to large (e.g.,
in filaments—76.9-168.1 by 25-75 wm).
According to den Hartog (1980), these
nematocysts are the most voluminous an-
thozoan cnidae, reaching up to 250 by 80
jxm. We also observed a particular shape of
the holotrichs I in the stomodaeum. They
varied from oblong to cylindrical (Figs. 41;
7G).
Holotrichs II (Figs. 3E, 6J, Table 1).—
Sharply distinguished from holotrich I in
capsule shape and in having small spines
with little contrast. The capsule is opaque,
cylindrical, with coiled tubule in small
tums, filling the entire undischarged cap-
sule. The spines are smaller than those of
holotrichs I and of little contrast and diffi-
cult to view with light microscopy.
Remarks.—These were exclusive to the
marginal tentacles in both species. An un-
usual variety of this category is smaller and
cylindrical, with a tubule filling the whole
capsule and with spines smaller than those
of the typical holotrich II. These capsules
seem to differ from the holotrich II, except
for their shape, with spines so reduced that
they appear like spots distributed inside the
capsule with irregularly coiled small turns.
Presently, we consider it a holotrich II, oc-
curring in the discal tentacles and in the
column of D. carlgreni (Figs. 2K 4E); in
the column of D. sanctithomae, they were
large (Fig. 7C). This variety has never been
reported from the discal tentacles and col-
umn in the Discosomatidae.
Discussion
We identified seven types of cnidae in the
Discosomatidae. The presence of spirocysts
in this group was discussed by Carlgren
(1949) and den Hartog (1980). Carlgren
(1949) recorded the sparseness of spirocysts
in the Discosomatidae, claiming that they
were found in the tentacles. den Hartog
(1980:36) characterized the family by the
absence of spirocysts and stated “‘not only
are spirocysts absent in the tentacles of Dis-
cosomatidae, other ectodermal cnidae too
are) very “scarse” .. ithe “tentacles: . <2 are
non-retractile, non-motile ... are either re-
VOLUME 113, NUMBER 1
WF
D. carlgreni ‘ ‘
a)
A
o
Cc
B
DISCAL
TENTACLE
‘“N
x
Oks
7
eens
Fig. 2. Cnidome of Discosoma carlgreni, Discal tentacles. A, b-rhabdoids?; B, b-rhabdoids'; C, D, p-rhab-
doids D'; E, G, holotrichs I; E holotrichs If. Abbreviation: t = terminal tubule. Scale: 10 wm.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
D. carlgrent
MARGINAL
TENTACLE
=>
SLL.
CLE —
\
SSA a5
y—
\
NAN
NAN
SS ANS AARNE
SSSA SD
Ce Sa ae a
LAT Or ire
N
~
~— =
Nes
, b-rhabdoids'!:
D, p-rhabdoids D'; E, holotrichs II; EK G, holotrichs I. Abbreviation: t = terminal tubule. Scale: 10 wm.
2.
’
Cnidome of Discosoma carlgreni, Marginal tentacles. A, spirocysts; B, b-rhabdoids
Bipg3)
a
om
VOLUME 113, NUMBER 1
D. carlgreni
COLUMN
STOMODAEUM
Cnidome of Discosoma carlgreni, Column A—F; Stomodaeum G—
: B, b-rhabdoids':
5
I. A, b-rhabdoids
Fig. 4.
C, D, G, p-rhabdoids D';
10 ym.
= terminal tubule. Scale:
E, holotrichs II; E H, I, holotrichs I. Abbreviation: t
?
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
2
FILAMENTS
D. carlgreni
| SN
~ Se oe
eee
te
ANS
<<
SE TNS
!
!
NA
=a At
=
Ww,
FW swans ay LIES pep OEE
Tea77 70! ‘\ wns Ss
rhabdoids D!. Abbreviation:
p
>
Cnidome of Discosoma carlgreni, Filaments. A, D, holotrichs I; B, C
t = terminal tubule. Scale: 10 wm.
Figs:
137
VOLUME 113, NUMBER 1
D. sanctithomae
MARGINAL
TENTACLE
DISCAL
Ww
—_/
<
Ee
z=
Lu
be
i eae ae ie ae I I
G, H,
B,
terminal
>
Fig. 6. Cnidome of Discosoma sanctithomae, Discal tentacles A-—D, Marginal tentacles E-K. A
p-rhabdoids D';
C, D, I, K, holotrichs I; E, spirocysts; EK b-rhabdoids'; J, holotrichs Il. Abbreviation: t
tubule. Scale: 10 xm.
°
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
3
D. sanctithomae
COLUMN
Sp PPE SS SSIS
> ss
we We %, “y Dy, “7
TTT,
SS Spam re
\
ol!
OTN
STOMODAEUM
Cnidome of Discosoma sanctithomae, Column A—F, Stomodaeum G-I. A, H, p-rhabdoids D!; B, b-
Fig. 7:
rhabdoids!;
10 pm.
terminal tubule. Scale:
—G, I, holotrichs I. Abbreviation: t
2.
>
holotrichs II; D, p-rhabdoids D
°
C
139
VOLUME 113, NUMBER 1
D. sanctithomae
~
FILAMENTS
SU iy FS 9
“Bh in Wee re
Z SLE
Iam
=
SS
Dopp 7Y
SSS >
GOD COCA a
=>
~
—_—
pee
POSAANA SASS
ML
-rhabdoids D!; C, D, holotrichs I. Abbre-
Cnidome of Discosoma sanctithomae, Filaments. A, B, p
Fig. 8.
viation: t
terminal tubule. Scale: 10 ym.
140
Table 1.—Distribution measurements of cnidae. Sp = species, A = Discosoma carlgreni, B = D. sanctith-
omae. n = number of capsules measured, F = figure.
Structure
Average and range of length
and width of nematocyst capsules
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Cnidae type Sp Length Width n jm
Discal tentacles
b-rhabdoids' A 14.4 (12.5-17.5) 4.7 (1.9-6.3) 41 2B
b-rhabdoids? A O57 .5=1 1-3) 1.8 (1.3-3.0) 41 2A
p-rhabdoids D'! A 12.9 (8.8-17.5) 3.5 (1.9-5.6) 66 ae
25.4 (18.8-30.6) 6.6 (5.6-8.8) 54 2D
B 13.3 (10.6—17) 3.5 (2.5-5.0) 40 6A
33.2 (20-46.9) 7.7 (4.4-11.3) 50 6B
holotrichs I A 37.2 (28-41.3) 13.3 (6.9-18.1) 76 2E
72.1 (45—123.1) 31.2 (18.1-55.6) 45 2G
B 34 (26.9-43.8) 14.8 (11.3-—20) 83 6C
69 (57.5-85) 32.4 (21.9-45.8) 92 6D
holotrichs I A 16.5 (11.3-—21.9) 4.9 (3.1-6.9) 42 2F
Marginal tentacles
b-rhabdoids' A 15.5 (12.2—23.8) 4.9 (3.8-6.9) 65 6) &
B 15.6 (11.3—20) 4.0 (2.5—5.6) 72 6F
b-rhabdoids? A 8.9 (6.3-10.9) 1.9 (0.9-3.1) 40 3B
p-rhabdoids D'! A 13.2 (8.8-16.3) 4.2 (1.9-6.9) 79 3D
B 13.5 (10-20) 3.8 (2.5—5.6) 50 6G
33.7 (23.1-49) 8.0 (5.6—11.9) 45 6H
holotrichs I A 35.6 (27.5—40.6) 12.7 (9.6—20.6) 65 3F
82.2 (50.6—125) 34.2 (20.6—-49.4) 60 3G
B 31.6 (25—34.4) 13.6 (9.4—-18.1) a2 6!
68.0 (54.4—-83.1) 30.0 (17.5—-40.3) 77 6K
holotrichs I A 29.7 (8.8-43.8) 5.4 (2.5—10.6) 159 3E
B 24.8 (13.1-—37) 5.1 (3.1-13.8) 110 6J
Column
b-rhabdoids'! A 15.7 (13.1—20) 5.5 (3.8-6.3) 63 4B
B 19.6 (16.3-25) 5.5 (3.8-7.5) 50 7B
b-rhabdoids? A 9.6 (5.6—12.5) 2.0 (1.3-3.8) 40 4A
p-rhabdoids D! A 13.9 (10-17.5) 4.6 (2.5-5.3) 60 4C
24.1 (19.1—28.8) 6.9 (6.2—8.1) 40 4D
B 14.5 (10—20.6) 4.0 (2.5-6.3) 45 TA
p-rhabdoids D? B 41.1 (25-58.8) 8.2 (4.4-13.1) 64 7D
holotrichs I A 34.6 (30.6—37.5) 1257 (73=16.3) 50 4F
B 34.1 (28.1—38.1) 15.4 (11.3-18.8) 58 TE
76.1 (61.9-94.4) 34 (25-42.5) 53 TF
holotrichs II A 20 (13.8-19) 6.3 (3.8—9.4) 65 4E
B 33.7 (26.3-40.6) 4.0 (2.5-6.3) 40 Ze
Stomodaeum
p-rhabdoids D' A 13.5 (8.8-19) 3.9 (1.9-6.3) 50 4G
B 17.9 (10.6—32.5) 4.7 (2.5—9.4) 52 7H
holotrichs I A 37.3 (32.5—40.6) 14.1 (10.6—17.5) 35 4H
54.6 (45.6—70) 18.1 (10.9—22.5) 65 4I
B 33.7 (25.640) 13.3 (5.6—18.8) a7 7
58.8 (44.4-78.8) 18.9 (11.9-35.6) 65 7G
Filaments
p-rhabdoids D'! A 18.2 (11.9-23.7) 4.9 (4.5-8.8) 63 5B
27 (25-33.1) 6.9 (5.0—10) 79 SC
B 14.6 (8.8—20) 3.6 (1.9-6.3) 40 8A
31.5 (24.4-38.1) 8.2 (5.0—-15) 78 8B
VOLUME 113, NUMBER 1 14]
Table 1.—Continued.
Average and range of length
and width of nematocyst capsules
Structure
Cnidae type Sp Length Width n F
holotrichs I A 36.9 (26.2—53.8) 13.6 (7.5—-18.8) 50 SA
117.6 (90.6—175.2) 49 (31.2-72) 69 5D
B 31.8 (23.8—37.5) 13.1 (5.0-18.1) 69 8C
146.5 (76.9—168.1) 60.3 (25-75) 73 8D
duced to insignificant, wartlike protuber-
ances or developed into vesicle-like struc-
tures...” den Hartog (1980) pointed out
the difficulty of accepting that these tenta-
cles were functional catching devices be-
cause the Discosomatidade have been as-
sociated with zooxanthellae that provide
nutrition to the animals. However, several
Actiniaria, such as Stichodactyla haddoni
(Saville-Kent, 1893) and Stichodactyla
duerdeni (Carlgren, 1900), with a large
number of spirocysts, have similar tentacles
and zooxanthellae. In this study, we found
spirocysts in the marginal tentacles of both
species, thus confirming Carlgren’s finding.
Although den Hartog (1980) accepted
Schmidt’s system (1972, 1974) of termi-
nology for corallimorpharian cnidae, he
preferred to adopt Stephenson’s system of
classification. den Hartog (1980) considered
Schmidt’s b- and p-rhabdoids (=b and p-
mastigophores sensu Carlgren, 1940) as
synonyms of spirulae and penicilli sensu
Stephenson (1928), respectively. Therefore,
he recorded four types of cnidae for the
Discosomatidae: spirulae, penicilli D, pen-
icilli E and homotrich. In agreement with
Belém & Schlenz (1982), we adopted
Schmidt’s classification because it provides
descriptions and illustrations of the differ-
ent types of corallimorpharian nematocysts,
based on many specimens.
The comparative study carried out with
specimens of Discosoma carlgreni from
Mexico and D. sanctithomae from Cuba has
confirmed that they are conspecific with the
Brazilian species, respectively.
The varieties of b-rhabdoids' and b-rhab-
doids? qualitatively separate Discosoma
carlgreni from the D. sanctithomae. The
first variety occurred particularly in the ten-
tacles as well as in the column of both spe-
cies, except in the discal tentacles of D.
sanctithomae. Such absence was also veri-
fied by den Hartog (1980). The b-rhab-
doids? occurred particularly in the tentacles
and column of D. carlgreni. Schlenz & Be-
lém (1982) recorded two size-classes of b-
rhabdoids in the stomodaeum of D. carl-
greni, neither of them observed by den Har-
tog (1980) nor in this study.
The p-rhabdoids D' were easily seen in
all structures of the body. The distribution
and occurrence of these nematocysts
seemed uniform in both species, except for
the presence of only one small size class in
the marginal tentacles of D. carlgreni as
well as in the column of D. sanctithomae.
Contrary to the view of den Hartog
(1980) concerning the total absence of p-
rhabdoid D in the stomodaeum in Coralli-
morpharia, we observed a variety p-rhab-
doid D' in this structure in both species.
These nematocysts were rather common,
also being larger in Discosoma sanctitho-
mae. Our results are in agreement with
those of Corréa (1964), who observed the
microbasic p-mastigophore (=p-rhabdoid D
sensu Schmidt) in the stomodaeum of D.
sanctithomae. We also observed the variety
p-rhabdoid D? in the column of D. sanctith-
omae. None of the previous works have
registered this variety before.
Another feature which distinguishes Dis-
cosoma sanctithomae from D. carlgreni is
the size of holotrich I. In the column of D.
sanctithomae, both large and small ones oc-
cur, whereas in D. carlgreni they are small.
142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
We noticed in all discharged capsules of
holotrich I, an abruptly tapered distal tube
called terminal tube by den Hartog (1980).
This terminal portion is flat, hardly refrac-
tive and spineless, being also observed by
Schlenz & Belém (1982). This portion can
be observed in squash preparations, though
sometimes it is not fully discharged so the
terminal tube of most remains unevaginat-
ed. The presence of this tube was previous-
ly recognized in the macrobasic p-masti-
gophores [=holotrich sensu Schmidt (1974)
and penicilli E sensu den Hartog (1980)] by
Cutress (1955:134). In his words: ‘“‘these
nematocysts have a shaft which is more
than three times the length of the capsule
and which is abruptly reduced to a
thread .. .”’. Cutress (1955), therefore, sug-
gested that corallimorpharian holotrichs
should be termed macrobasic p-mastigo-
phores. Nevertheless, some authors (e.g.,
Werner 1965, Mariscal 1974) did not accept
Cutress’ proposal. An alternative and in-
dependently derived system, using Stephen-
son’s term penicilli E, was provided by den
Hartog (1980). Schmidt (1972, 1974) iden-
tified a large holotrich I, commonly occur-
ring in the filaments, stomodaeum and ten-
tacles in the Corallimorpharia, as well as in
the Scleractinia, as having the most distinc-
tive spines possessed by anthozoan nema-
tocysts. As indicated by Schmidt (1972,
1974), this type presented a gradual taper-
ing of the tubule.
Studying the cnidae of four species of
Brazilian Mussidae, Pires & Pitombo
(1992) observed a holotrich I in the mes-
enterial filaments. However, they did not re-
cord in this type the abrupt end of the tube
as a vestigial thread. In short, Pires & Pi-
tombo (1992) observed a gradual tapering
of the tubule, as mentioned by Schmidt
(1974).
den Hartog et al. (1993), who studied the
corallimorpharians from the CANCAP ex-
pedition, found penicilli E, especially in the
filaments, as well as in the tentacles, of five
species of Corallimorphidae. However, they
did not observe the terminal tube in dis-
charged capsules of penicilli E from the fil-
aments of Corynactis sp. den Hartog et al.
(1993) added that the previous observations
on this type by Cutress (1955) and espe-
cially by den Hartog (1980) needed confir-
mation, considering the fact that the peni-
cilli E were based on few occasions.
In spite of the divergence among termi-
nologies adopted in previous works, we still
consider the type holotrich I sensu Schmidt
(1972, 1974) the best term to be employed
in this paper. To avoid further misunder-
standing, the holotrichs I found in Disco-
somatidae here has an abrupt tapered and
spineless distal tip into a terminal tubule
(the reason that den Hartog (1980) consid-
ered them as penicilli E).
The holotrichs II of the marginal tenta-
cles of Discosoma carlgreni are in variably
larger than those of D. sanctithomae. This
nematocyst was also found by den Hartog
(1980). According to den Hartog (1980)
and Belém & Schlenz (1982), this type oc-
curs only in the marginal tentacles. Never-
theless, an unusual variety of holotrich, ob-
served in the column and discal tentacles,
showed size differences, especially in the
column of D. sanctithomae. This variety
had never been found before. We tentative-
ly classify it as Schmidt’s holotrich II; fur-
ther studies of its ultrastructure will provide
a more comprehensive description.
The results of this study demonstrate the
importance of the nematocysts in distin-
guishing Discosoma carlgreni from D.
sanctithomae. The varieties of b-rhabdoids
and p-rhabdoids D allow us to separate the
species.
Acknowledgments
We are grateful to Dr. E. Schlenz and Dr.
E L. da Silveira (Universidade de Sao Pau-
lo, Sao Paulo) for their helpful comments
to the manuscript. Thanks to Dr. A. Herrera
(Instituto de Oceanologia, Academia de
Ciencias de Cuba, ACC), Dr. E. Jordan
(Universidade Nacional Autonoma de Méx-
ico), Dr. E M. Amaral (Universidade Fed-
VOLUME 113, NUMBER 1
eral Rural de Pernambuco) and Dr. FE B.
Pitombo (Universidade Federal Rural do
Rio de Janeiro) for providing some of the
samples used in this study. We also owe
thanks to Dr. E Pitombo for the photograph
of Figure 2B. Thanks to Parque Nacional
Marinhos dos Abrolhos who provided fa-
cilities and assistance during the collection
of specimens. We wish to thank Conselho
Nacional de Desenvolvimento Cientifico e
Tecnol6gico (CNPq), Brazil for financial
support. Part of this study was also sup-
ported by FAPESP (Fundacgao de Amparo
a Pesquisa do Estado de Sao Paulo), by a
grant to the first author (no 98/3222-2). Fi-
nally, we also grateful to Dr. Daphne Fautin
(University of Kansas) and an anonymous
reviewer who made invaluable suggestions
that improved the manuscript.
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Appendix
List of specimens from the Museu Nacional do Rio
de Janeiro Cnidaria collection used for this study.
Discosoma carlgreni (Watzl, 1922)
MNRJ. 1796—Brazil, Espirito, Santo Santa Cruz
(19°49'08"S and 40°16'43”W), Aracruz, Estagao de
Biologia Marinha, coll. S. M. Pinto, E B. Pitombo
& EM. Amaral, 8 Aug 1990, 14 specimens, det. S.
M. Pinto Aug 1990.
MNRJ. 1866 and 1867—Brazil, Espirito Santo, Santa
Cruz (19°49'08"S and 40°16'43”"W), Aracruz, Esta-
¢ao de Biologia Marinha, coll. M. J. C. Belém & E.
Schlenz, 8 Sep 1991, 14 specimens, det. M. J. C.
Belém & E. Schlenz Sep 1991.
MNRJ. 1868—Brazil, Espirito Santo, Guarapari
(20°40'16”S and 40°28’5”W), Trés Praias, coll. M. J.
C. Belém & E. Schlenz, 5 Sep 1991; 1 specimen,
det. S. M. Pinto Sep 1991.
MNRJ. 1878—Brazil, Espirito Santo, Santa Cruz
(19°49'08"S and 40°16'43”W), Aracruz, Estagao de
Biologia Marinha, coll. M. J. C. Belém, E. Schlenz
& C. C. Ratto, 8 Sep 1991, 1 specimen, det: M. J.
C. Belém Aug 1991.
MNRJ. 2075—Brazil, Bahia, Abrolhos Archipelago
(17°20’—18°10'S and 38°35’—39°20'W), Siriba Island,
coll. E B. Pitombo & C. G. Fonseca, 15 Dec 1992,
5 specimens, det. S. M. Pinto Dec 1992.
MNRJ. 2250—Brazil, Bahia, Abrolhos Archipelago
(17°20'—18°10'S and 38°35’—39°20'W), Siriba Island,
coll. FE B. Pitombo,& C. C., Ratto, 19 Dec. 199358
specimens, det. S. M. Pinto Dec 1993.
MNRJ. 2259—Brazil, Bahia, Abrolhos Archipelago
(17°20’-18°10'S and 38°35’—39°20'W), Chapeirao,
coll. E B. Pitombo, 23 Dec 1993, 13 specimens, det.
S. M. Pinto Dec 1993.
MNRJ. 1540—México, Puerto Morelos, Quintana
Roo, coll. E. Jordan Dahlgren & E D. Amaral, 24
Oct 1989, 1 specimen, det: S: M- Pinto Dec 1989:
Discosoma sanctithomae (Duchassaing & Michelotti,
1860)
MNRJ. 2076—Brazil, Bahia, Abrolhos Archipelago
(17°20'—18°10'S and 38°35’—39°20'W), Siriba island,
coll. EK B. Pitombo & C. G. Fonseca, 15 Dec 1992,
22 specimens, det. S. M. Pinto Dec 1992.
MNRJ. 2251 and 2252—Brazil, Bahia, Abrolhos Ar-
chipelago (17°20’—18°10’S and 38°35’—39°20'W),
Siriba island, coll. EK B. Pitombo & C. C. Ratto, 19
Dec 1993, 20 specimens, det. S. M. Pinto Dee 1993.
MNRJ. 2109—Cuba, La Habana, Playa, Playa Jaiman-
itas, coll. A. Herrera, 26 Aug 1992, 4 specimens, det.
M. J. C. Belém Feb 1994.
MNRJ. 2110—Cuba, Recife de Punta del Este, Isla de
Juventud, coll. A Herrera, 26 Aug 1992, 4 speci-
mens, det. M. J. C. Belém Dec “1992.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):145—154. 2000.
Additions to the cancellariid (Mollusca: Neogastropoda) fauna of
South Africa
Richard E. Petit and M. G. Harasewych
Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560-0118, U.S.A.
Abstract.—Four new species of Cancellariidae are described from the con-
tinental shelf and upper continental slope off eastern South Africa. Admetula
afra is distinguished from all congeners by its combination of small adult size
(<11 mm), rounded shoulder, evenly reticulate sculpture, and distinct varix at
the juncture of protoconch and teleoconch. Trigonostoma kilburni differs from
all other Trigonostoma in its distinctive shell outline, unornamented peripheral
keel, rounded rather than tabulate shoulder, and very narrow umbilicus. Nip-
ponaphera wallacei differs from N. paucicostata (Sowerby, 1894), its geo-
graphically closest congener from the Arabian Sea area, in being umbilicate
and in having a more rounded shoulder, and more numerous and finer spiral
cords. Its frequent association with the turbinid Bolma andersoni suggests that
it may be an ectoparasite of this species. Zeadmete verheckeni is most similar
to Zeadmete subantarctica Powell, 1933, from off New Zealand, from which
it can be distinguished by its lower spire and weaker surface sculpture. Among
the South African taxa, Z. verheckeni most closely resembles ‘‘Cancellaria”’
eutrios Barnard, 1959, from which it is easily distinguished by its tabulate
shoulder and lower spire. The geographic and bathymetric ranges of Admetula
epula Petit & Harasewych, 1991, a species previously known only from ‘“‘ex
pisces”’ material, have been expanded based on live-collected specimens.
The Cancellariidae comprises a family of
diverse and highly specialized, suctorial
neogastropods that inhabit soft bottom, sub-
tidal to bathyal habitats throughout tropical
and temperate seas. The cancellariid fauna
of South Africa was reviewed comprehen-
sively by Barnard (1959) and Kensley
(1973), and to a limited extent more re-
cently in popular works by Richards (1981)
and Steyn & Lussi (1998).
This paper describes four new species of
cancellariids collected in South African wa-
ters by SCUBA and by the vessels R/V
Meiring Naudé (1984-1988) and NMPD
Africana (1995). These species are assigned
to the genera Admetula, Trigonostoma, Nip-
ponaphera, and Zeadmete. The new taxa
are compared to related species from the
Indian Ocean, New Zealand, and Australia.
All type specimens are housed in the col-
lections of the Natal Museum (NM), Pie-
termaritzburg, Republic of South Africa.
Family Cancellariidae Forbes & Hanley,
1851
Genus Admetula Cossmann, 1889
Type species: Cancellaria evulsa (Solan-
der, 1766) (=Buccinum evulsum Solander,
1766) by original designation.
Admetula epula Petit & Harasewych, 1991
Figs. 1-2, 18
Admetula epula Petit & Harasewych,
1991:181, figs. 1-3.
Diagnosis.—A small species with an
ovately conical shell. Transition from pro-
146 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
toconch to teleoconch gradual, indicated by
appearance first of spiral, then axial sculp-
ture. Teleoconch sculpture of strong spiral
cords and less pronounced, rounded axial
ribs. Outer lip thin, smooth within.
Gross anatomy.—Preserved animal yel-
lowish tan, foot long, narrow, tapering pos-
teriorly. Mantle cavity spanning 0.67 whorl.
Osphradium slightly broader than ctenid-
ium. Pericardium very small. Tentacles
symmetrical, bluntly cylindrical, flanking
central rostrum. Eyes small, black. Probos-
cis short, about 0.67 length of mantle cav-
ity. Buccal mass large, nearly filling retract-
ed proboscis. Salivary glands and accessory
salivary glands in cephalic haemocoel, not
contained within proboscis. Penis long, nar-
row, dorsoventrally compressed, distal end
bluntly rounded.
Remarks.—This species was originally
described based on seven specimens taken
from the stomachs of fish. Its bathymetric
range was inferred to overlap with that of
Congiopodus spinifer (Smith) (55—146 m),
one of the fish from which it was taken.
Among the material collected by the
NMPD Africana [sta. A17419D] was a live
collected specimen of A. epula (Figs. 1-2,
18) trawled at a depth of 210 m off the
mouth of the Tsitsikamma River.
We have examined a specimen of Ad-
metula from deeper water (450—500 m) off
the Mbashe River, Transkei (R/V Meiring
Naudé sta. Q14), between the type localities
of A. epula and the new species. This spec-
imen (NM C9050) is too worn for the tran-
sition from protoconch to teleoconch to be
clearly discerned, but there is no indication
of a pronounced varix. Because it also has
secondary spiral cords between all primary
cords, this specimen is tentatively identified
as A. epula. This record expands the geo-
graphic range of A. epula from Cape St.
Blaize to Transkei, off of the mouth of the
Mbashe River (32°22.8’S, 29°00.8’E). The
bathymetric range is extended well into the
bathyal zone (450 m).
Admetula afra, new species
Figs. 3—4, 18
Diagnosis.—A small species with a
broadly conical shell. Protoconch demar-
cated from teleoconch by broad, rounded
varix. Teleoconch with sharply reticulated
sculpture. Outer lip reflected posteriorly,
with weak lirae beneath spiral cords.
Description.—Shell (Fig. 3) small, reach-
ing 9.0 mm, ovately conical with rounded
anterior. Protoconch (Fig. 4) of 1.67
smooth, inflated whorls deflected from coil-
ing axis of teleoconch by about 7°. Transi-
tion to teleoconch (Fig. 4, arrow) demar-
cated by prominently rounded varix, fol-
lowed immediately by onset of both spiral
and axial sculpture. Teleoconch of 2.75—3
evenly rounded whorls. Suture weakly im-
pressed. Shoulder rounded, weakly defined.
Axial sculpture consists of narrow, regular-
ly spaced, weakly prosocline ribs (14-16
on body whorl). Spiral sculpture of narrow,
sharply defined primary cords (12-13 on
body whorl, 6 on penultimate whorl). Sin-
gle, weak, secondary cords may be present
between primary cords on posterior half of
body whorl. Spiral and axial sculpture in-
tersect to form sharply reticulated pattern
with small nodes at intersections of ribs and
cords. Aperture broadly ovate, deflected
from coiling axis by 16—19°. Outer lip thin,
slightly flared posteriorly, weakly lirate be-
neath spiral cords. Parietal callus thin,
translucent, overlying 5—6 spiral cords from
previous whorl. Columella forming angle of
128-130° with parietal region, thick,
straight, with two weak columellar folds
and broad siphonal fold. Siphonal canal
shallow but well-defined. Shell color white.
Type locality.—SE of Port Durnford,
South Africa (29°01.5'S, 32°11.8’E), dredged
in 310-320 m, glutinous sandy mud. R/V
Meiring Naudé sta. ZQ9, 6 Jul 1985.
Type material.—Holotype, NM_ E3189/
T1382, 8.0 mm; Paratype 1, 7.8 mm, and
Paratype 2, 6.9 mm, NM V7006/T1383 from
the type locality; Paratype 3, 9.0 mm, NM
E3764, from off Cape Vidal (28°08.4'S,
VOLUME 113, NUMBER |
Figs. 1-4. 1-2; Admetula epula Petit & Harasewych, 1991. Off Tsitsikamma River, South Africa (34°45'S,
24°47'E) in 210 m, sand, old shell grit and shell debris. NMPD Africana sta. A17419D. 1. Apertural view of
shell. 2. Apical view of protoconch. 3—4. Admetula afra new species. Holotype, NM E3189/T1382, SE of Port
Durnford, South Africa (29°01.5’S, 32°11.8’'E), dredged in 310—320 m, glutinous sandy mud. R/V Meiring Naudé
sta. ZQ9, 6 Jul 1985. 3. Apertural view of holotype. 4. Apical view of protoconch.
32°36.4'E), dredged in 165 m, moderately Comparative remarks.—This new spe-
fine sand, R/V Meiring Naudé sta. ZM8, 11 cies appears most closely related to the
June 1988. more southern Admetula epula Petit & Har-
Etymology.—Feminine form of the Latin asewych, 1991, from which it differs in
afer, African. having uniformly rectangular sculpture
148 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
formed by equally-sized axial ribs and spi-
ral cords, and a posteriorly reflected outer
lip with weak denticles beneath the spiral
cords. The most striking difference between
these two species is the presence of a strong
varix at the termination of the protoconch
in A. afra. In contrast, the transition from
protoconch to teleoconch is indistinct and
gradual in Admetula epula.
Genus Trigonostoma Blainville, 1827
Type species: Delphinula trigonostoma
Lamarck, 1822 (?=Buccinum scalare Gme-
lin, 1791) by monotypy.
Trigonostoma kilburni, new species
Figs. 5-10, 18
Diagnosis.—A small species with a thin,
angular, narrowly umbilicate shell. Shoul-
der rounded, not tabulate, lacking pro-
nounced spines. Axial sculpture of numer-
ous scabrous varices. Outer lip smooth,
lacking lirae.
Description.—Shell (Fig. 5) small, reach-
ing 14.3 mm, thin, angular, biconical,
strongly shouldered, with deep, narrow um-
bilicus. Spire high (spire angle 55°), com-
prising over half of shell length. Protoconch
(Figs. 6—7) of 2 smooth whorls, offset from
coiling axis of shell by about 5°. Transition
to teleoconch abrupt, marked by a slightly
flared varix and the onset of spiral sculp-
ture. Teleoconch of up to 5 sharply angular
whorls. Suture deeply impressed behind the
evenly rounded shoulder delineated by a
sharp keel along the periphery of the shell.
Axial sculpture consists of numerous,
flared, weakly prosocline varices (Fig. 8),
regularly spaced on early whorls (about 16—
18 on first teleoconch whorl), increasing in
number and becoming more irregularly
spaced in subsequent whorls. Spiral sculp-
ture of broad, crisply demarcated primary
spiral cords (Fig. 8, p), with 1-3 slightly
narrower secondary cords (Fig. 8, s) be-
tween adjacent primary cords, and much
finer spiral threads (Fig. 8, t) between some
cords. Aperture roundly triangular, deflect-
ed from coiling axis by 18—19°. Siphonal
canal short, broad, barely discernible except
externally as the siphonal fasciole. Outer lip
thin, smooth within, slightly reflected with
spiral cords visible through edge of lip.
Posterior portion of inner lip adpressed
against siphonal fasciole. Short parietal re-
gion forms angle of 150° with long, slightly
concave columella that bears 2 weak, wide-
ly spaced columellar folds and 1 siphonal
fold. Umbilicus deep, narrow. Shell white,
sometimes with a yellowish cast.
Radular teeth (Figs. 9-10) extremely
long, ribbon-like, tricuspid. Central cusp
smooth, with recurved rim. Lateral cusps
long, folded toward central cusp, each with
four, anteriorly directed secondary cusps.
Second most proximal secondary cusp bi-
fid.
Type locality.—Off East London, South
Africa (33°04.7'S, 28°07.2’E) dredged in 90
m, associated with coarse sand, sponges,
gorgonians. R/V Meiring Naudé sta. XX46,
17 Jul 1984.
Type material.—Holotype, NM D679,
12.9 mm, from type locality. Paratype, NM
D680, 14.4 mm, off Kidd’s Beach, South
Africa (33°11.8'S, 28°03.2’E) dredged in 90
m, associated with coarse sand, sponges. R/
V Meiring Naudé sta. XX50, 17 Jul 1984.
Etymology.—This species honors Dr. Ri-
chard N. Kilburn, Natal Museum, Pieter-
maritzburg, Republic of South Africa, for
his many contributions to malacology.
Comparative remarks.—This new spe-
cies differs from all other Trigonostoma in
its distinctive shell outline. It may be dis-
tinguished from Trigonostoma scalare
(Gmelin, 1791) and T. thysthlon (Petit &
Harasewych, 1987) by its unornamented
peripheral keel, its rounded rather than tab-
ulate shoulder, and by its very narrow um-
bilicus. The only other South African Tri-
gonostoma is the common shallow-water T.
semidisjuncta (Sowerby, 1849), which has
a heavier, more rounded shell with strong
spiral cords.
VOLUME 113, NUMBER I
149
Figs. 5-8.
(33°04.7'S, 28°07.2'E), dredged in 90 m, coarse sand, sponges, gorgonians. R/V Meiring Naudé sta. XX46, 17
Jul 1984. 5. Apertural and lateral views of holotype. 6. Apical and 7. Lateral views of protoconch. 8. Detail of
surface sculpture on body whorl. p, primary spiral cords; s, secondary spiral cords; t, spiral threads.
Genus Nipponaphera Habe, 1961
Type species: Nipponaphera habei Petit,
1972 by I.C.Z.N. Opinion 1052.
Nipponaphera wallacei new species
Figs. 11-13, 16
Diagnosis.—A small species with a
heavy, strongly sculptured, narrowly um-
bilicate shell. Aperture sharply triangular,
outer lip with strong teeth along inner edge
Trigonostoma kilburni new species. Holotype, NM D679, Off east London, South Africa
of varix. Columella with two prominent,
Sharply keeled columellar folds and a
strongly reflected siphonal fold.
Description.—Shell (Fig. 11) small for
genus, to 12.4 mm, ovately conical, with
rounded anterior. Spire relatively short
(spire angle 70°), comprising less than half
of shell length. Protoconch (Fig. 13) of 1.67
smooth, inflated whorls. Transition to teleo-
conch marked by onset of spiral cords fol-
lowed immediately by strong axial ribs. Te-
150
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs. 9-10. Trigonostoma kilburni new species. Distal ends of radular teeth of holotype. 9. Lateral view.
10. End-on view.
leoconch of up to 4 angular whorls. Suture
impressed behind sloping shoulder that is
delineated by peripheral keel. Axial sculp-
ture of strong, broad, prosocline ribs (10 on
penultimate whorl) that become more wide-
ly spaced on body whorl where they appear
as varices with flared edges. Spiral sculp-
ture of broad, flattish primary cords, 3 sec-
ondary cords between adjacent primary
cords, with fine threads between some
cords. Aperture sharply triangular, deflected
from coiling axis by 20°. Siphonal canal
short, well-defined. Outer lip reflected an-
teriorly, with spiral sculpture visible
through thin edge, 9 strong teeth along in-
ner edge of varix, small, angular indenta-
tion at shoulder. Parietal region short, col-
umella straight, with small, thin callus, 2
Sharply keeled columellar folds and strong-
ly reflected siphonal fold. Umbilicus nar-
row, inconspicuous, bordered by well-de-
veloped, cord-like siphonal fasciole. Shell
color chestnut brown, with lighter spiral
bands along shoulder and middle of body
whorl.
Type locality.—Off Phumula, Natal,
South Africa, in 45 m, on reef. Jun 1997.
Type material.—Holotype, NM V4689,
6.6 mm, from type locality. Paratype 1, NM
V3889, Off Phumula, Natal, South Africa,
in 35 m, taken by SCUBA, living on shell
of Bolma andersoni (E. A. Smith, 1902). 4
Sep 1996. Paratype 2, NM V303, 12.4 mm,
Off Park Rynie, Natal, South Africa, in 53
m. Taken by SCUBA, living on the spire of
the turbinid gastropod Bolma andersoni. 19
May 1990.
Etymology.—This species is named for
Mr. Martin Wallace, who collected the type
material and generously made it available
for study.
Comparative remarks.—The genus Nip-
ponaphera is distinguished from Trigonos-
toma primarily by the presence, in the latter,
of an open umbilicus that extends back to
the protoconch. Also, Nipponaphera has a
wide and flat siphonal fold. Although the
new species here described has an umbili-
cus, it is not profound. It differs from N.
paucicostata (Sowerby, 1894) of the Ara-
bian Sea area in being umbilicate, in having
a more rounded shoulder, and more and fin-
er spiral cords. Specimens of N. paucicos-
tata figured by Verhecken (1986, Figs. 4—
6) lack varix-like axial ribs on the body
whorl, but the type specimen has very
strong, unevenly spaced ribs on the body
whorl. The Japanese N. teramachii (Habe,
1961) has a sharp keel, is umbilicate, with
a cord-like siphonal fasciole, but lacks the
squarish, finely imbricated spiral sculpture
of this new species. Melvill and Standen
(1901:451) reported N. paucicostata from
the Gulf of Aden ‘adhering to the upper
VOLUME 113, NUMBER 1
Figs. 11-13.
SCUBA in 45 m, on reef. Jun 1997. 11. Apertural view of holotype. 12. Detail of sculpture on body whorl. 13.
Apical view of protoconch.
part of Rapana bulbosa, 30—50 fathoms.”
The alimentary system of cancellariids is
adapted to feed on body fluids of prey/host
organisms (Petit & Harasewych, 1986; Har-
asewych & Petit, 1986). The association of
species of Nipponaphera with other gastro-
pods suggests that this group of cancellar-
lids may be specialized ectoparasites of
large gastropods.
Genus Zeadmete Finlay, 1926
Type species: Cancellaria trailli Hutton,
1973 by original designation.
Zeadmete verheckeni, new species
Figs. 14-18
Diagnosis.—A small species with an
ovate shell. Shoulder tabulate. Surface
sculpture dominated by spiral cords. Pseu-
do-umbilicus narrow. Outer lip thin, smooth
within. Buccal mass minute. Radula absent.
Description.—Shell (Fig. 14) small,
reaching 7.9 mm, thin, with stepped spire,
rounded anterior. Protoconch (Figs. 15—16)
erect, smooth, of 1.5 whorls. Transition to
Nipponaphera wallacei new species. Holotype, NM V4689, Off Phumula, Natal, South Africa,
teleoconch abrupt, marked by onset of
weak, closely-spaced axial ribs that become
stronger, more regularly spaced, when spi-
ral cords first appear within 0.25 whorl. Te-
leoconch of 2 rounded, strongly tabulate
whorls. Suture strongly impressed. Axial
sculpture of evenly spaced, well-defined,
axially aligned ribs (30 on body whorl) as
broad as intervening spaces. Spiral sculp-
ture of sharply demarcated evenly spaced
cords (2 between suture and shoulder, 17
below shoulder) diminishing in strength
abapically, forming small nodules as they
cross axial ribs. Aperture elongated, nar-
rowly elliptical. Outer lip faintly sinuate,
smooth within. Columella with two broad,
weak, columellar folds (Fig. 17, pcf, acf)
and siphonal fold (Fig. 17, sf). Siphonal ca-
nal small but distinct, axially aligned, not
forming siphonal fasciole. Anterior portion
of inductura bordered by weak parietal
wash, partially covering narrow pseudo-
umbilicus. Shell color white. Periostracum
thin, finely lamellate, straw-colored.
Preserved animal white, with short, nar-
row, posteriorly rounded foot. Tentacles tu-
[52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
1 mm
sf
500 um
“
og ge
3 a
_
Figs. 14-17.
Zeadmete verheckeni new species. Holotype, NM C6800, Off Stony Point, Transkei, South
Africa (32°37.5'S, 28°45.8’E), dredged in 390—400 m, muddy sand, small stones. R/V Meiring Naudé sta. V11,
12 Jul 1984. 14. Apertural and dorsal views of holotype. 15. Apical and 16. Lateral views of protoconch. 17.
Columella. pcf, posterior columellar fold; acf, anterior columellar fold; sf, siphonal fold.
bular, symmetrical, with very large black
eyes at their bases. Penis long, narrow, dor-
so-ventrally flattened, with small terminal
papilla. Osphradium very broad, strongly
asymmetrical, dorsal leaflets twice as broad
as ventral leaflets. Ctenidium less than half
as wide and twice as long as osphradium.
Hypobranchial gland large, glandular. Re-
tracted proboscis occupies anterior two-
thirds of cephalic haemocoel, the rest oc-
cupied by long, convoluted mid-esophagus.
Proboscis strongly coiled within proboscis
sheath. Extended proboscis likely exceeds
shell length. Proboscis thin, with minute
buccal mass in anteriormost 0.125 of pro-
boscis. Radula absent.
Type locality.—Off Stony Point, Trans-
kei, South Africa (32°37.5’S, 28°45.8’E),
VOLUME 113, NUMBER 1
015° E
020° E
Fig. 18.
025° E
ee
030° E 035° E
Geographic distributions of South African Cancellariidae. Admetula epula Petit & Harasewych,
1991, vertically hatched area = type locality; open squares = new records. Admetula afra new species, star in
circle = type locality; filled circle = additional record. Trigonostoma kilburni new species, black star = type
locality; open circle = additional record. Nipponaphera wallacei new species, triangle = type locality. Zeadmete
verheckeni new species, diamond = type locality.
dredged in 390—400 m, muddy sand, small
stones. R/V Meiring Naudé sta. V11, 12 Jul
1984.
Type material.—Holotype, NM C6800,
5.1 mm, from type locality.
Etymology.—Named for Mr. André Ver-
hecken, Mortsel, Belgium, in recognition of
his contributions to the study of the Can-
cellariidae.
Comparative remarks.—Placement of
this new species in the genus Zeadmete is
tentative. Zeadmete verheckeni is concho-
logically most similar to a group of species
from off the southern coasts of New Zea-
land and Australia that have been assigned
to the genera Oamaruia Finlay, 1924 and
Zeadmete Finlay, 1926. Powell (1979:224)
treated Zeadmete as a subgenus of Oama-
ruia. Several New Zealand and Australian
species presently assigned to Zeadmete
agree with this South African species in
Shell form and sculpture (see Garrard 1975,
Powell 1979). Zeadmete subantarctica
Powell, 1933, from 50 fathoms off Snares
Islands, New Zealand, has the same shell
shape and columellar structure as Z. ver-
heckeni but has deeply cancellated sculp-
ture on the posterior half of the body whorl
and only spiral cords on the anterior por-
tion.
In the course of a study on Australian
cancellariids, we found that the abyssal
Zeadmete kulanda Garrard, 1975 has a rad-
ula similar to that of Nothoadmete tumida
(Oliver, 1982:figs. 3, 5). The fact that no
radula was found in Z. verheckeni argues
against these species being congeneric.
However, we are reluctant to introduce an-
other genus-level taxon until more data on
the species with this shell form are avail-
able. The problems of generic placement of
small, deep-water cancellariids was briefly
discussed by Verhecken (1997:296).
Among the South African taxa, this spe-
cies most closely resembles ‘‘Cancellar-
ia’ eutrios Barnard, 1959, from which it is
154 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
easily distinguished by its tabulate shoulder
and lower spire.
Acknowledgments
We thank Dr. Richard N. Kilburn for
making these specimens available and for
his patience in waiting for their description.
Correspondence and discussion with Mr.
André Verhecken of Mortsel, Belgium add-
ed to our knowledge and was helpful in pre-
paring this paper.
Literature Cited
Barnard, K. H. 1959. Contributions to the knowledge
of South African marine Mollusca. Part II. Gas-
tropoda: Prosobranchiata: Rachiglossa.—An-
nals of the South African Museum 45:1—237.
Garrard, T. A. 1975. A revision of Australian Cancel-
lariidae (Gastropoda: Mollusca).—Records of
the Australian Museum 30(1):1—62.
Harasewych, M. G., & R. E. Petit. 1986. Notes on the
morphology of Admete viridula (Gastropoda:
Cancellariidae).—The Nautilus 100(3):85—91.
L.C.Z.N. 1976. Opinion 1052. Nipponaphera Habe,
1961 (Gastropoda): Designation of a type-spe-
cies under the plenary powers.—Bulletin of
Zoological Nomenclature 32(4):242—243.
Kensley, B. 1973. Sea-Shells of southern Africa, Gas-
tropods. Maskew Miller Ltd., Cape Town, 236
PP.
Melvill, J. C., & R. Standen. 1901. The Mollusca of
the Persian Gulf, Gulf of Oman, and Arabian
Sea, as evidenced mainly through the collec-
tions of Mr. EF W. Townsend, 1893-1900; with
descriptions of new species.—Proceedings of
the Zoological Society of London for 1901:
327-460, pls. 21-24.
Oliver, P. G. 1982. A new species of cancellariid gas-
tropod from Antarctica with a description of the
radula.—British Antarctic Survey Bulletin 57:
15-20.
Petit, R. E., & M. G. Harasewych. 1986. New Philip-
pine Cancellariidae (Gastropoda: Cancellari-
acea), with notes on the fine structure and func-
tion of the Nematoglossan radula.—The Veliger
28(4):436-443.
Powell, A. W. B. 1979. New Zealand Mollusca. Wil-
liam Collins, Auckland, xiv + 500 pp.
Richards, D. 1981. South African seashells. A collec-
tor’s guide. C. Struik Publishers, Cape Town. 98
pp. + 60 pls.
Steyn, D. G., & M. Lussi. 1998. Marine shells of South
Africa. Ekogilde Publishers, Hartebeespoort.
264 pp.
Verhecken, A. 1986. A revision of the Cancellariidae
(Neogastropoda: Cancellariacea) of the Red Sea
and the Gulf of Aden.—Gloria Maris 25(4):
133-153.
Verhecken, A. 1997. Mollusca Gastropoda: Arafura
Sea Cancellariidae collected during the KA-
RUBAR Cruise. Pp. 295-323 in A Crosnier &
P. Bouchet, eds., Résultats des Campagnes MU-
SORSTOM, vol. 16. Mémoirs du Muséum Na-
tional d’ Histoire Naturelle 172.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):155—-161. 2000.
Erpobdella lahontana (Annelida: Hirudinea: Arhynchobdellida:
Erpobdellidae), a new species of freshwater leech from
North America
Peter Hovingh and Donald J. Klemm
(PH) 721 Second Avenue, Salt Lake City, Utah 84103, U.S.A.;
(DJK) U.S. Environmental Protection Agency, National Exposure Research Laboratory,
Ecological Exposure Research Division, Ecosystems Research Branch,
26 W. Martin Luther King Drive, Cincinnati, Ohio 45268, U.S.A.
Abstract.—new species of a leech, Erpobdella lahontana, is described from
the Lahontan Basin in California and Nevada of the western United States.
This species has four pairs of eyes, the preatrial loops of male paired ducts
extend to ganglion XI, and the male and female gonopores are located in
furrows of the annuli, separated by five annuli.
The zoological history of the Great Basin
of the western United States has resulted in
an area of high endemism of fishes (Hubbs
& Miller 1948) and hydrobiid snails
(Hershler 1998). However, amphibians have
not morphologically evolved into endemics
and may be the result of very recent im-
migration to the region (Hovingh 1997).
A general survey of the Great Basin and
adjacent regions of the western United
States was undertaken to determine if
leeches had drainage specific distributions.
Over 2300 aquatic sites were surveyed in
the Great Basin including some 600 sites
within the Lahontan Basin. Leeches show
an entirely different pattern than fishes, hy-
drobiid snails, and amphibians with respect
to basin specific distribution without mor-
phological evolution, suggesting evolution-
ary stasis and an inhabitant of the Great Ba-
sin since its geological formation in the
Miocene (Hovingh, unpublished data).
Within this study, an erpobdellid leech was
identified with the gonopores separated by
five annuli. This leech was found in two
separate drainages of the Honey Lake Sub-
basin (Eagle Lake and one other location
out of the 2300 sites in the survey) in the
Lahontan Basin of northeastern California.
This paper describes this new leech species
and is the first endemic species of leech to
be found in the Great Basin.
Materials and Methods
Collection methods consisted of exam-
ining the underside of substrates such as
rocks, logs, and anthropogenic debris in the
periphery of aquatic systems (i.e., springs,
streams, and lakes) up to 100 cm deep.
Leeches were relaxed with dilute ethanol,
wiped clean of mucous, fixed in 10% for-
malin overnight, and preserved in 70% eth-
anol. Histological examination of serial sec-
tions of the clitellum region, after staining
with hematoxylin and eosin, was used for
detailed morphological analysis of the gen-
ital atrium, cornua, and preatrial loops of
the ejaculatory ducts. A model of the gen-
ital atrium, cornua, and preatrial loops was
constructed from photographs taken of the
serial sections.
Systematics
Family Erpobdellidae Blanchard, 1894
Genus Erpobdella Blainville, 1918
Erpobdella lahontana, new species
Type material.—Holotype, United States
National Museum (USNM 186409) and 8
156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
paratypes (USNM 186410), deposited in
the National Museum of Natural History,
Division of Worms, Smithsonian Institu-
tion, Washington, D.C., collected 15 Sep
1997, same locality.
Type locality.—Eagle Lake, Lassen
County, California; latitude 40°33.5’N and
longitude 120°48.8’W (selected by the
abundance of leeches at this locality and
public access to the lake).
Diagnosis.—Dorsal surface dark gray,
heavily mottled with light gray spots; white
or yellow irregular minute, transverse rows
of papillae on the dorsal side of every an-
nulus with some living specimens having
prominent papillae on both dorsal and ven-
tral sides; dorsum with a light black mid-
dorsal stripe or plain; five-annulate, each
annulus of approximately equal size; four
pairs of eyes: first pair of labial eyes large,
second pair of smaller labial eyes behind
first pair; buccal eyes two pairs, small; male
and female gonopores separated by five an-
nuli, located in furrows of segments XII
and XIII and male gonopore large, raised,
and especially glandular (Fig. 1); atrium
with cornua (horns), with sperm ducts
forming paired preatrial loops extending an-
teriorly to ganglion XI (Figs. 2, 3). Preatrial
loops angle laterally and anteriorly from
cornua. Cornua and preatrial loops dip ven-
trally to join each other. Ovisacs extend
posteriorly to ganglion XV. Five annuli sep-
arate the segmental ganglia.
Description of Holotype
External anatomy.—(based on holotype,
USNM 186409): Body elongate, flattened,
sides of body narrowing along most of
length to pointed head (length 30 mm, max-
imum width 3 mm); body wall uniformly
smoke-gray, dorsal surface darker gray than
ventral surface, dorsal surface with a faint
middorsal dark line, no black pigmentation;
dorsal surface dark gray, heavily mottled
with light gray spots; somites five-annulate;
white irregular minute, transverse rows of
papillae on every annulus; clitellum con-
<— 5mm —P
Fig. 1. Erpobdella lahontana. Ventral view, male
and female gonopores (holotype).
spicuous, 15 annulate; nephridiopores on b,
annulus of XI and XII (other nephridio-
phores not determined); mouth small (width
of oral sucker 1 mm); eyes four pair, one
large labial pair, second pair of small labial
eyes behind first pair and two smaller buc-
cal pairs on fourth annulus (further poste-
rior); anus surrounded by papillae, located
dorsally at the base of the caudal sucker;
caudal sucker small (width 2 mm), less than
one-half maximum body width; male gon-
opore large, raised, cylindrical and glandu-
lar, (Fig. 1) located in furrow of the clitel-
lum region between annuli XIIb, and XIIb,;
distance from mouth to male gonopore, 14
mm; female gonopore inconspicuous, lo-
cated in furrow between annuli XIIIb, and
XIIb,.
Internal anatomy.—(based on dissection
of paratypes): Atrium wider than long; atri-
al horns projecting anteriorly; preatrial
loops of vas deferens extending to ganglion
XI; ovisacs extend from segments XIII to
XV.
Additional observations of paratypes
(USNM_ 186410).—External anatomy.
Paratypes resemble the holotype with the
following additional observations: length
VOLUME 113, NUMBER 1
157
Fig. 2.
Ganglion XI (denoted A) and Ganglion XII (denoted D) showing the preatrial loops (denoted B) and the genital
atrium (denoted C) with cornua. The angle in which the preatrial loops join the cornua is indicated by short
bars: 30° for the left side and 45° for the right side as viewed from the dorsum. Note that the right cornua
displays an oval cross-section representing the section that has turned ventrally. Long bar = | mm.
(42—55 mm) and width (3.5—5 mm) of some
individuals varies from the holotype; clitel-
lum is inconspicuous in some individuals;
female gonopore difficult to locate in some
individuals; anus small to large in some
paratypes.
Remarks.—The eyes in four pairs are ar-
ranged in two transverse rows; mouth with
muscular ridges but no jaws; body seg-
ments (somites) five-annulate with all an-
nuli of approximately equal in width. The
structure of the reproductive system shows
that this species belongs to the genus Er-
pobdella Blainville, 1918, with preatrial
loops of the male paired ducts and testes in
small, numerous bunched sacs.
Distribution, habitat, and ecology.—Er-
pobdella lahontana is found in two subba-
sins of the western Lahontan Basin (Fig. 4):
Eagle Lake and Grasshopper Valley in Las-
Photograph of horizontal (parallel to dorsal and ventral surfaces) section of the region between
sen County, California. The Pleistocene
precursor of Eagle Lake and Grasshopper
Valley (Madeline Plains) drained into Hon-
ey Lake Subbasin, the latter being an arm
of Pleistocene Lake Lahontan. The new
species was found in Buck Bay in the north
part of Eagle Lake and along the south
shore of Eagle Lake. In Eagle Lake, the
leech was often found on vesicular basaltic
rocks. When removing the leech, the pos-
terior sucker would remain on the rocks.
This occurred with five leeches after which
sampling was from smoother artificial sub-
strate. The leech was found in the outflow
of springs in Grasshopper Valley. Eleva-
tions varied from 1555 to 1625 m above
mean sea level (msl). Eagle Lake and, to a
lesser extent, the Grasshopper Valley were
associated with ponderosa pine (Pinus pon-
derosa) forest and both belong to the Mo-
158
Fig: 3.
ejaculatory ducts. Legend same as in Figure 2. Draw-
ing made from a model constructed from photographs
of serial sections of this region. The arrows denote
where the cornua (paired horns) and the preatrial loops
both turn ventrally before they are joined. The right
cornua also makes a turn to the center.
Dorsal view of male genital atrium and
doc Plateau geomorphic province, a region
of numerous basalt-andesite flows which
isolated these two basins from Honey Lake
(Gester 1962). Eagle Lake (12,150 ha) has
three subbasins of which two (north and
central) basins do not stratify due to their
Shallow nature and strong winds. Conduc-
tivity of Eagle Lake is near 800 pMbhos,
probably as a result of the fact that 40% of
Eagle Lake’s inflow is derived from ground
water (Huntsinger & Maslin 1976). Eagle
Lake contains five species and Madeline
Plains contains one species of native fish.
The ecology of Eagle Lake is further de-
scribed by Huntsinger & Maslin (1976).
Two other locations in the Lahontan Basin
(Winnemucca and Granite Springs Subba-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
sins, east of Pyramid Lake, Nevada) con-
tained leeches in which the gonopores were
separated by five annuli, and these popula-
tions are still under investigation.
Associated leech species.—Erpobdella
lahontana was associated with the leeches
Erpobdella punctata (Leidy, 1870), Helob-
della stagnalis (Linnaeus, 1758), and Hae-
mopis marmorata (Say, 1824). In Grass-
hopper Valley, H. marmorata was associ-
ated with E. lahontana. Eagle Lake speci-
mens at the National Museum of Natural
History, Worm Division, include: E. punc-
tata (USNM 42502, USNM 60061, USNM
60062), Glossiphonia complanata (USNM
2572), Placobdella ornata (USNM 60055),
Theromyzon trizonare (USNM 42500), and
H. marmorata (USNM 42570). The leech
specimen (USNM 42571) was classified as
Dina fervida, but this specimen is most
likely E. lahontana. The gonopore separa-
tion was not unresolved, but the pigmenta-
tion and number of eyes suggested this spe-
cies.
Variations.—All Eagle Lake specimens
(22) were identical with respect to eyes and
the number of annuli between gonopores.
Some specimens contained dark annuli in
which the papillae became conspicuous
with an even darker dorsal strip, and this
varied to uniform light color with invisible
papillae and with no dorsal strip. Sizes of
leech specimens were up to 55 mm long
and 3.5 mm wide.
Similar species.—Erpobdella lahontana
is distinguished from E. dubia (Moore &
Meyer, 1951), E. parva (Moore, 1912), and
E. punctata (Leidy, 1870) by the five annuli
separating the gonopores. Erpobdella dubia
(dorsum greenish, heavily mottled, usually
with a variable dark mid-dorsal stripe; gon-
opores separated by 3.5—4 annuli) and E.
parva (dorsum unpigmented or color uni-
formly smokey gray; gonopores separated
by 2.5—3.5 annuli) have not been observed
in the Great Basin collections with the gon-
opores separated by five annuli. If the gon-
opores are not distinguishable, the pigmen-
tation patterns of E. lahontana could lead
VOLUME 113, NUMBER 1 159
LL a a a a a a a LF SBT IES ae: |
Lassen 5
Madeline Plains
ho”
Fig. 4. Map of the western Lahontan Basin in California and Nevada. Lassen County is enclosed and the
communities of Susanville, California and Reno, Nevada are designated by diamonds. The heavy solid line
denotes the Lahontan Basin with arrows showing Pleistocene drainages from Eagle Lake and Madeline Plains
Subbasins. The light solid line shows the extent of Pleistocene lakes with the patterned horizontal lines showing
the present Eagle, Honey, and Pyramid Lakes. The three solid circles show location of Erpobdella lahontana in
Eagle Lake and in Grasshopper Valley of the Madeline Plains Subbasin. Inset shows the locations in relation to
the Great Basin and the western United States. Map derived from Snyder et al. (1964).
160
to some confusion with E. dubia and E.
parva, neither of which has been found in
the Lahontan Basin. E. punctata has been
found in the Great Basin, but this species
can be distinguished easily from E. lahon-
tana by having only two annuli separating
the male and female gonopores, and having
three pairs of eyes. Taxonomic discussion
and keys to the family Erpobdellidae and
these North American species can be found
in Sawyer (1972, 1986b), Klemm (1985,
1990, 1995) and Davies (1991).
Discussion
Similar species, Erpobdella dubia and E.
parva, to the new species, E. lahontana,
have been historically classified within the
genus Dina (Soos 1963, 1966, 1968; Saw-
yer 1972, Klemm 1985,,Davies 1991):-The
genus (or subgenus) Dina is identified by
the fact that every fifth annulus (b,) of a
somite is distinctly wider or more broad
than the other four annuli and is subdivided
by a faint transverse furrow (Lukin 1976,
Mann 1982, Sawyer 1986a, 1986b; Davies
1991, Nesemann 1995, Neubert & Nese-
mann 1995). Also, the genus Dina is re-
ported from only the southern, western, and
central areas of the Palaearctic region (Lu-
kin 1976, Sawyer 1986b). Therefore, Saw-
yer (1986a, 1986b) revised the genus Er-
pobdella to include the two species of Dina
from the Neartic region, including North
America, based on the equal width of all
the body annuli and distribution of E. dubia
and E. parva. It was concluded that the ge-
nus Dina was not found in North America.
This was later confirmed by Klemm (1990,
1995). The North America (Neartic region)
genera Nephelopsis, Mooreobdella, and the
Palearctic region genus Dina were grouped
together by the feature that, of any five an-
nuli of the body somites, there is one an-
nulus that is distinctly wider than the other
four annuli and usually subdivided by a
faint transverse furrow (Mann 1962, Lukin
1976, Sawyer 1986b, Davies 1991). This
anatomical feature was not seen in Great
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Basin species of Erpobdellidae. All the spe-
cies of erpobdellids in North America have
annuli of similar width.
Etymology.—Louis Armand de Lom
d’ Arce, Baron de Lahontan was a 17th cen-
tury French explorer of the upper Missis-
sippi River basin who suggested that the
western continental flows drained into a sa-
line lake and then into the ocean. Although
Baron de Lahontan may never have entered
the western drainages or the Great Basin,
the large western basin, Lahontan Basin,
within the Great Basin was named after the
Baron (Cline 1963). The leech is, thus,
named after the basin in which it is found.
The anatomical tradeoff in erpobdellid
leeches having preatrial loops and having
the gonopores separated by more annuli is
that the preatrial loops and the cornua may
become shorter. An evolutionary compen-
sation would be for the cornua and the pre-
atrial loop to form contortions. This may be
what is happening in Erpobdella lahontana
with the cornua and the preatrial loops both
taking a ventral dip to their junction.
Acknowledgments
The authors wish to thank Teresa Puste-
jovsky (U.S.ES., Eagle Lake, Susanville,
California) for a wealth of ecological infor-
mation and Cathy Sanderson, Wasatch His-
to Consultants, Winnemucca, Nevada for
making the histological slides of the repro-
ductive system.
Literature Cited
Cline, G. C. 1963. Exploring the Great Basin.—Uni-
versity of Oklahoma Press, Norman, Oklahoma,
254 pp.
Davies, R. W. 1991. Annelida: leeches, polychaetes,
and acanthobdellids. Pp. 427-479 in J. H.
Thorp and A. P. Covich, eds. Ecology and clas-
sification of North American freshwater inver-
tebrates. Academic Press, Inc., New York, 911
Pp.
Gester, G. C. 1962. The geological history of Eagle
Lake, Lassen County, California.—Occasional
Papers of California Academy of Sciences 34:
1-29.
Hershler, R. 1998. A systematic review of the hydro-
VOLUME 113, NUMBER 1
biid snails (Gastropoda: Rissoida) of Great Ba-
sin, western United States. Part 1. Genus Pyr-
gulopsis.—Veliger 41:1—132.
Hovingh, P. 1997. Amphibians in the eastern Great Ba-
sin (Nevada and Utah, USA): a geographical
study with paleozoological models and conser-
vation implications.—Herpetological Natural
History 5(2):97-134.
Hubbs, C. L., & R. R. Miller. 1948. The zoological
evidence. The Great Basin, with emphasis on
glacial and post-glacial times.—Bulletin Uni-
versity of Utah Biological Series 38:18—166.
Huntsinger, K. R., & P. E. Maslin. 1976. A limnolog-
ical comparison of the three basins of Eagle
Lake, California.—California Fish and Game
62:232-245.
Klemm, D. J. 1985. Freshwater leeches (Annelida: Hir-
udinea). Pp. 70-194 in D. J. Klemm, ed., A
guide to the freshwater Annelida (Polychaeta,
naidid and tubificid Oligochaeta, and Hirudinea)
of North America. Kendall/Hunt Publishing
Company, Dubuque, Iowa, 198 pp.
. 1990. Hirudinea. Pp. 398—415 in B. L. Peck-
arsky, P. R. Fraissinet, M. A. Penton, & D. J.
Conklin Jr. eds. Freshwater macroinvertebrates
of northeastern North America. Cornell Univer-
sity Press, Ithaca, New York, 442 pp.
. 1995. Identification guide to the freshwater
leeches (Annelida: Hirudinea) of Florida and
other southern states.—Bureau of Surface Water
Management, Florida Department of Environ-
mental Protection, 2600 Blair Stone Road, Tal-
lahassee, Florida 32399-2400.
Lukin, E. I. 1976. Fauna of the USSR. Leeches, vol.
1. Leeches of fresh and brackish water bod-
ies._—Academy of Science, Zoological Institute
of the Soviet Union. Nauka Publishers, Lenin-
grad Branch. (467 pp., Translated from Russian
by Literature Research Company for the U.S.
EPA (in part) and the Canadian Translation Bu-
16]
reau for the Canadian Museum of Nature (in
part).
Mann, K. H. 1962. Leeches (Hirudinea) their structure,
physiology, ecology, and embryology. Perga-
mon Press, New York, 201 pp.
Nesemann, H. 1995. On the morphology and taxono-
my of the Asian leeches (Hirudinea: Erpobdel-
lidae, Salifidae)—Acta Zoologica Academiae
Hungaricae 41:165—182.
Neubert, E., & H. Nesemann. 1995. Contribution to
the knowledge of the genus Dina Blanchard,
1892 (Hirudinea: Erpobdellidae).—Hydrobio-
logia 315:89-94.
Sawyer, R. T. 1972. North American freshwater leech-
es, exclusive of the Piscicolidae with a key to
all species.—Illinois Biological Monographs
46:1-154.
. 1986a. Leech Biology and Behavior. volume
I. Anatomy, physiology, and behavior. Oxford
University Press, Oxford, 417 pp.
. 1986b. Leech Biology and Behavior. volume
II. Feeding Biology, Ecology, and Systematics.
Oxford University Press, pp. 419-793.
Snyder, C. T., G. Hardman, & F. F Zdenek. 1964. Pleis-
tocene Lakes in the Great Basin. U.S.G.S. Mis-
cellaneous Geological Investigations Map _ I-
416.
Soos, A. 1963. Identification key to the species of the
genus Dina R. Blanchard, 1892 (Emend. Mann
1952, Proceedings Zoological Society, London
122) (Hirudinea: Erpobdellidae).—Acta Uni-
versitatis Szedgediensis, Szeged, Hungaricae
IX:253-261.
. 1966. Identification key to the leech (Hirudi-
noidea) genera of the world, with a catalogue
of the species. III. Family Erpobdellidae.—Acta
Zoologica Academiae Scientiarum Hungaricae
XII. (3—4):37 1-407.
. 1968. Identification key to the species of the
genus Erpobdella de Blainville, 1818 (Hirudi-
noidea: Erpobdellidae).—Annales Historico-
Naturales Musei Nationalis Hungaricae 60:141—
145.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):162—209. 2000.
A cladistic analysis of Sciomyzidae Fallén (Diptera)
Luciane Marinoni and Wayne N. Mathis
(LM) Department of Zoology, Universidade Federal do Parana, Caixa Postal 19020, 81531-990,
Curitiba, Parana, Brazil;
(WNM) Department of Entomology, NHB 169, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.
Abstract.—A preliminary cladistic analysis of adult characters is presented
that illustrates the phylogenetic relationships among the genera of the family
Sciomyzidae. The monophyly of Sciomyzidae is based primarily on larval char-
acters: the habit of malacophagy and the presence of a serrate ventral arch that
articulates with the lower margin of the mouth hooks. A reduction in the num-
ber of spermathecae, from three to two, is also likely to be a synapomorphy
for Sciomyzidae. The analysis was done using Hennig&6, and 37 morphological
characters were arranged among 50 genera. After using successive weighting,
six cladograms were produced, and from these a consensus cladogram was
obtained. The subfamilies Salticellinae and Sciomyzinae are confirmed to be
monophyletic, as are the tribes Sciomyzini and Tetanocerini. Eutrichomelina
Steyskal, which has been placed in the tribe Sciomyzini, is transferred to the
tribe Tetanocerini. The genus Antichaeta Haliday is confirmed to be in the tribe
Tetanocerini. Illustrations of antenna and male terminalia are presented.
Among families of Acalyptrate Diptera,
the Sciomyzidae Fallén (1820), more com-
monly known as marsh or snail-killing flies,
are comparatively well studied, especially
the biology of their immature stages (Berg
& Knutson 1978, Ferrar 1987, Knutson
1987, Rozkosny 1997). Sciomyzid larvae
are primarily parasitoids or predators on
aquatic or terrestrial mollusks. This feeding
proclivity may be of considerable impor-
tance to the biological control of certain
parasitic, mostly tropical diseases, such as
fascioliasis and schistosomiasis (Knutson
1976). The trematodes causing both diseas-
es parasitize many of the same aquatic mol-
lusks, as intermediate hosts, that are also
fed upon by sciomyzid larvae.
Although the natural history and ecology
and to a degree the descriptive taxonomy
and cytology (Boyes et al. 1972) of the
Sciomyzidae are relatively well known, no
cladistic analysis at the generic level is
available. As a step toward filling that void
in our knowledge, this cladistic study was
undertaken and is reported here. To provide
perspective, we begin this report with a
brief overview of the higher-level classifi-
cation.
In the first comprehensive treatment of
Palearctic Sciomyzidae, Hendel (1900) di-
vided the family into two subfamilies: Scio-
myzinae and Tetanocerinae. Hendel char-
acterized these subfamilies by the proepi-
sternal seta (present in Sciomyzinae, absent
in Tetanocerinae) and the frontal vitta (well
developed and shiny in most Tetanocerinae,
absent or reduced in most Sciomyzinae).
Cresson (1920), in a study limited to the
Nearctic fauna of Sciomyzidae, recognized
Sciomyzinae, as characterized by Hendel,
and described two additional subfamilies:
Dryomyzinae and Euthycerinae. Cresson
also proposed five tribes that were divided
among two of the subfamilies as follows:
(1) Sciomyzinae with Oidematopsini and
Sciomyzini; and (2) Euthycerinae with
VOLUME 113, NUMBER |
Chaetomacerini, Euthycerini, and Sepedon-
tini.
Hendel (1924) published a key to the Pa-
learctic genera and distinguished the genus
Tetanura Fallén as a separate subfamily,
Tetanurinae, based on the following com-
bination of characters: arista subapical,
forefemur bare, and ovipositor telescoped.
In the same paper and within the subfamily
Tetanocerinae, Hendel also proposed the
tribe Salticellini for the genus Salticella Ro-
bineau-Desvoidy.
Over 30 years ago, Steyskal (1965) pro-
posed a classification for the Sciomyzidae
that has been the most extensive treatment
for the family from the standpoint of higher
categories. Steyskal’s classification, which
has been adopted by most subsequent work-
ers (Knutson et al. 1976, Barnes 1979, Roz-
kosny & Elberg 1984, Knutson 1987,
Barnes & Knutson 1989, McAlpine 1989),
recognized five subfamilies: Huttonininae,
Salticellinae, Helosciomyzinae, Phaeomyi-
inae, and Sciomyzinae. The subfamily Scio-
myzinae included two tribes, Sciomyzini
and Tetanocerini. Although Steyskal’s char-
acterization of Sciomyzidae did not identify
apomorphic characters, he distinguished the
family from related families by the follow-
ing set of morphological characters: costal
vein (C) without breaks; subcostal vein (Sc)
complete, free from vein R,; vein A, com-
plete; oral vibrissae absent; postvertical se-
tae divergent to parallel; midfemur bearing
a seta on the anterior surface; and at least
one tibia with a preapical seta (Knutson
1987, McAlpine 1989). The tribes Tetano-
cerini and Sciomyzini are distinguished by
the presence (Sciomyzini) or absence (Te-
tanocerini) of a proepisternal seta.
Griffiths (1972), who incorporated many
characters of the male genitalia in his high-
er level phylogenetic study, introduced the
prefamily as a category between the super-
family and the family categories. Using this
classificatory structure, the prefamily Scio-
myzoinea comprises the families Coelopi-
dae, Phaeomyiidae, Dryomyzidae, Scio-
myzidae, Helosciomyzidae, Ropalomeridae,
163
Sepsidae, Megamerinidae, and Cremifani-
dae. Griffiths’ studies of Sciomyzidae were
based on the following species: Pherbellia
guadrata Steyskal, Pherbellia griseola
(Fallén), Sciomyza simplex Fallén, Pterom-
icra apicata (Loew), Elgiva sundewalli
Kloet & Hincks, and Tetanocera robusta
(Loew). The family Sciomyzidae, as char-
acterized by Griffiths (1972), includes Sal-
ticellinae + Sciomyzinae, and the family’s
monophyly is based primarily on the ma-
lacophagous habits of the larvae and the
presence of a ventral arch in the cephalo-
pharyngeal skeleton (Knutson et al. 1970,
Barnes 1981, McAlpine 1989). The reduced
number of spermathecae, from three to two,
is likely to be another synapomorphy for
the Sciomyzidae, although in Salticellinae,
the number of spermathecae is four. We in-
terpret the latter condition to be secondarily
derived from two and to be an autapomor-
phy for the subfamily Salticellinae.
The subfamily Salticellinae has three
species in two genera, the extant Salticella
Robineau-Desvoidy and the fossil Prosal-
ticella Hennig. The Sciomyzinae, which in-
clude 505 recent species, have 57, mostly
widespread genera.
The purpose of this paper is to present a
classification for the genera of Sciomyzidae
sensu Griffiths that is based on a cladistic
analysis of primarily morphological char-
acters. Our analysis is intended more spe-
cifically to test the hypotheses that the sub-
families Salticellinae and Sciomyzinae and,
within the latter subfamily, the tribes Scio-
myzini and Tetanocerini are monophyletic.
Material and Methods
Fifty of the 57 genera belonging to the
Salticellinae and Sciomyzinae were exam-
ined and analyzed (Appendix 1). The seven
genera not examined are: Ditaeniella Sack,
Eulimnia Tonnoir & Malloch, Neodictya
Elberg, Oligolimnia Mayer, Pseudomelina
Malloch, Tetanoptera Verbeke, and Verbek-
aria Knutson. We also studied a new genus
and species from India, Steyskalina picta
164 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Ghorpadé & Marinoni, that was recently
described (Ghorpadé et al. 1999). Since it
was virtually impossible to examine all spe-
cies of each genus and suspecting that some
genera may be polyphyletic (e.g., Pherbel-
lia Robineau-Desvoidy with 81 species; Se-
pedon Latreille with 75 species; and Tetan-
ocera Duméril with 49 species) the analysis
was done with the type species serving as
exemplars of each genus. The type species
are always linked to the appropriate generic
name.
Pelidnoptera, represented by P. fuscipen-
nis (Meigen 1830), was used as the out-
group. This genus, which is in the family
Phaeomylidae (Griffiths 1972), was select-
ed because it is morphologically similar and
closely related, perhaps the sister group
(Steyskal 1965, included it as a subfamily)
of the Sciomyzidae. Its characters are di-
rectly comparable to those of Sciomyzidae.
The family Helosciomyzidae (Griffiths
1972), represented by Helosciomyza aliena
Malloch (1928); Huttonina abrupta Tonnoir
& Malloch (1928); Huttonina furcata Ton-
noir & Malloch (1928), and Huttonina scu-
tellaris Tonnoir & Malloch (1928), was
studied to further confirm the polarization
of characters.
A matrix with 37 morphological charac-
ters of adults and one of the larval mor-
phology was produced (Table 1). Autapo-
morphies for particular genera were not in-
cluded in the analysis.
Multistate characters in the analysis were
first treated as unordered (Carvalho 1989,
Pape 1992, Marinoni & Carvalho 1993).
The ordination of the characters was done
in accordance with standard procedures for
cladistic analysis (Wiley 1981). The polar-
ization was done using the outgroup com-
parison (Watrous & Wheeler 1981, Wiley
1981, Brooks 1989).
The analysis was facilitated with the
computer program Hennig86, version 1.5
(Farris 1988). To find cladograms supported
by the most consistent characters, the fol-
lowing command sequence was used:
‘“‘mhennig”’ (mh), “‘branch and swapping”
(*) and “‘successive weighting’ (xs w)
(Carpenter 1988, Dietrich & McKamey
1995). The option “‘nelsen’’ (ne) was used
to construct a strict, consensus cladogram
(Pape 1992, Marinoni & Carvalho 1993).
Characters and Character States Used in
the Analysis
The characters used in the analysis are
listed and discussed in the same sequence
as they appear in the cladogram (Figs. 323—
324). The letters A and P represent the rel-
ative apomorphic (derived) and plesio-
morphic (primitive) conditions respectively.
la. Larval feeding behavior: P (0) feed-
ing on Diplopoda; A (1) feeding as a par-
asitoid or predator on terrestrial or aquatic
Mollusca.
This is one of just a few characters in the
matrix that establishes the monophyly of
Salticellinae + Sciomyzinae. The genus
Pelidnoptera is a parasitoid on Diplopoda,
and numerous larval characters distinguish
it from the larvae of Sciomyzidae (Vala et
al. 1990). Although malacophagy charac-
terizes nearly all Sciomyzidae and is a syn-
apomorphy for the family, there is at least
one species, Sepedonella nana Verbeke,
that feeds on oligochaetes (Vala et al 2000)
as a secondary departure from the ground-
plan of the more basal clades of the family.
1b. Ventral arch in the larval cephalo-
pharyngeal skeleton: P (0) absent; A (1)
present.
The serrated ventral arch articulates with
the ventral margin of the larval mouth
hooks. This is the only larval structural
character that is an autapomorphy for Sal-
ticellinae + Sciomyzinae and may be relat-
ed with the malaphagous feeding behavior.
2. Number of spermathecae: P (0) three;
A (1) two; A (2) four.
According to McAlpine (1989) the im-
mediate ancestor of Muscomorpha had
three sclerotized spermathecae. This is the
basic number in the main sections of Mus-
comorpha: Aschiza, Schizophora, Acalyp-
tratae, and Calyptratae. In Pelidnoptera, the
VOLUME 113, NUMBER 1
Table 1.—Matrix of taxa and character states.
Pelidnoptera
Salticella
Sciomyza
Oidematops
Atrichomelina
Tetanura
Colobaea
Calliscia
Parectinocera
Pteromicra
Pherbellia
Eutrichomelina
Ectinocera
Renocera
Antichaeta
Chasmacryptum
Shannonia
Perilimnia
Hoplodictya
Dictya
Hydromya
Neolimnia
Tetanoceroides
Euthycerina
Tetanocera
Trypetolimnia
Psacadina
Steyskalina
Dictyodes
Ilione
Pherbina
Trypetoptera
Limnia
Poecilographa
Pherbecta
Protodictya
Guatemalia
Elgiva
Hedria
Dichetophora
Coremacera
Dictyacium
Euthycera
Ethiolimnia
Teutoniomya
Thecomyia
Sepedoninus
Sepedonella
Sepedon
Sepedomerus
Sepedonea
000
123
001
120
LLO
110
000
456
010
010
120
020
120
iy oP
110
v6
010
010
010
OEE
101
Od.
101
10,2
101
FOP
101
101
101
DOs
COL
102
G2
102
102
nO
Oe
101
00.
101
101
101
1.1
TOL
101
nO
FO
£04
101
£02
Jo
101
101
102
102
102
102
LZ
1 02
000
789
a
012
000
020
101
101
100
110
£10
000
000
000
000
000
000
000
020
000
000
000
000
000
000
000
010
000
000
010
000
000
010
010
010
ili BL
345
000
000
000
000
000
000
000
100
100
000
000
001
011
011
010
111
678
000
000
000
000
000
000
000
010
000
000
000
000
000
000
100
110
112
TAZ
ai 2
2
12
12
142.
142
eee
{ise
MEY
Es
cag
wi
ele
ce
aia
LAB
142
Ti2Z
ly
qb2
ited
dl:
ie i
101
a ba
003
a Se
pee
isl s
113
TVS
113
13
LZ
901
000
000
000
000
000
000
000
000
000
000
000
000
001
000
000
000
001
001
222
234
000
000
000
000
000
000
000
000
000
000
000
000
000
000
100
000
000
000
000
000
000
000
000
000
000
000
000
000
140
100
100
100
100
100
aaa ad
ie eh
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
222
567
000
000
000
000
000
010
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
000
165
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166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
number of spermathecae is also three, this
number being considered the plesiomorphic
condition for the Sciomyzidae. Only in Sal-
ticella is the number of spermathecae four,
which we interpret to be a secondary con-
dition, probably being derived from two.
Thus, four spermathecae are an autapomor-
phy for the Salticellinae, and two sperma-
thecae remain a synapomorphy for Scio-
myzinae.
3. Position of sixth left abdominal spi-
racle of the male (Figs. 115-159): P (O) in
membrane; A (1) in sclerotized tergite.
Within the Sciomyzidae the abdominal
spiracles of males may occur in the mem-
brane or the sclerotized tergite. Primitively
in Muscomorpha, the spiracles are in the
membrane, which is considered the ple-
siomorphic condition for the Sciomyzidae.
The position of the sixth spiracle in the
sclerotized portion of the tergite is a syna-
pomorphy for the Tetanocerini. Males of
Colobaea Zetterstedt have the spiracle in
the sclerotized tergite, a condition that
probably represents a secondary reversal.
4. Sixth abdominal tergite of the male
(Figs. 69, 74-76): P (O) present; A (1) ab-
sent.
Having all abdominal sclerites present is
undoubtedly the plesiomorphic condition.
According to McAlpine (1989) the reduc-
tion of the sixth abdominal tergite in males
is an apomorphy for the superfamily Scio-
myzoidea. Griffiths (1972) considered this
reduction to be a synapomorphy, confirm-
ing the monophyly of Sciomyzidae. In
Sciomyzidae, however, there is a complete
absence of this tergite in most genera. Only
in Salticella and in four genera of Scio-
myzini, Oidematops Cresson, Parectino-
cera Becker, Pherbellia, and Pteromicra
Lioy, is there a sixth tergite, which, how-
ever, is reduced.
5. Subepandrial plate (Figs. 161-169,
197): A (QO) absent; P (1) vestigial; A (2)
well developed.
The common, plesiomorphic condition,
is the presence of a vestigial subepandrial
plate found in Pelidnoptera, Salticella, Co-
lobaea, Calliscia Steyskal, Parectinocera,
Pteromicra, and Pherbellia. It is lacking in
all genera of the Tetanocerini. In Sciomyza
Fallén, Oidematops, and Atrichomelina
Cresson, the subepandrial plate is well de-
veloped and is a synapomorphic condition
that characterizes these three genera. In Te-
tanura (tribe Sciomyzini), the plate is also
absent, a condition we consider to be ho-
moplastic.
6. Anterior surstylus (Figs. 161—203): P
(O) well developed; A (1) vestigial; A (2)
absent.
The transformation series for this char-
acter is linear: 0 ~ 1 ~ 2. Pelidnoptera
has two pairs of well-developed, paired sur-
Styli, and thus the presence of an anterior
well-developed surstylus is the plesiom-
orphic condition for this character. Within
the Sciomyzidae, the tribe Sciomyzini also
has an anterior, well-developed surstylus.
The genera Tefanura (tribe Sciomyzini) and
Elgiva Meigen; the assemblage formed by
the genera Euthycerina Malloch, Tetano-
cera, Trypetolimnia Mayer, Psacadina En-
derlein, and Steyskalina Knutson; and the
group of Thecomyia Perty, Sepedoninus
Verbeke, Sepedonella Verbeke, Sepedon,
Sepedomerus Steyskal + Sepedonea Steys-
kal; lack an anterior surstylus, having only
the posterior one. The remaining genera of
Tetanocerini have a vestigial, anterior sur-
stylus.
7. Shape of aedeagus (Figs. 61—63, 66—
67, 204-322): P (O) asymmetrical; A (1)
symmetrical.
Asymmetry in the postabdomen of male
Acalyptratae may involve internal struc-
tures, including the aedeagus. Within gen-
era of Sciomyzidae, the aedeagus demon-
strates great variation in shape and sym-
metry. Pelidnoptera has an asymmetrical
aedeagus, a condition that is considered to
be plesiomorphic. In most Tetanocerini, ex-
cept for Antichaeta, the aedeagus is a com-
plex structure with several completely
asymmetrical sclerites. In the Sciomyzini
and in the group of Sepedoninus, Sepedo-
VOLUME 113, NUMBER 1
nella, Sepedon, Sepedomerus + Sepedonea,
the aedeagus is completely symmetrical.
8. Attachment of gonopod (Figs. 204—
322): P (O) fused to the hypandrium; A (1)
free.
McAlpine (1989) postulated the fusion of
the gonopod to the hypandrium as a ple-
siomorphic condition for the Acalyptratae.
All groups examined that are closely related
to the Sciomyzidae have the gonopod fused
to the hypandrium. The presence of a gon-
opod that is well developed and free is a
condition that supports the monophyly of
the tribe Sciomyzini. In Tetanoceroides
Malloch, a free gonopod is apparently ho-
moplastic.
9. Paramere (Figs. 204—322): A (QO) not
digitiform, elongate and well developed; P
(1) digitiform and well developed; A (2) ab-
sent.
Pelidnoptera has a digitiform, well-de-
veloped paramere that is considered to be
plesiomorphic. In Salticella, the paramere is
elongate and well developed, as in the Scio-
myzini and in the genera Antichaeta, Psa-
cadina, Neolimnia, Ethiolimnia, and Steys-
kalina. Two genera, Sepedonea and Sepe-
donella, jack a paramere, a condition that is
considered homoplastic.
10. Shape of aedeagal apex (Figs. 204—
213): P (O) variously shaped, but not flat-
tened; A (1) flattened.
The presence of an aedeagus with a flat-
tened apex is a synapomorphy for the group
of Sciomyza, Oidematops, Atrichomelina,
Tetanura, and Colobaea within the Scio-
myzini.
11. Basiphallus (Figs. 203-322): P (0)
well developed; A (1) present but poorly
developed; A (2) absent.
This character has the greatest number of
steps in the matrix and analysis, indicating
a large number of homoplasies and rever-
sals. The taxa that have the intermediate
apomorphic condition, the basiphallus
poorly developed, are Colobaea, Tetanura,
Trypetolimnia, Tetanoceroides, Dictyodes
Malloch, Jlione Verbeke, Pherbina Robi-
neau-Desvoidy, Trypetoptera Hendel, Gua-
167
temalia Steyskal, Hedria Steyskal, Diche-
tophora Rondani, Poecilographa Melander,
Protodictya Malloch + Pherbecta Steyskal
(the later two genera are sister groups, as
indicated by the “‘+”’’ connection). The gen-
era Salticella, Antichaeta, Sepedonella, Se-
pedon, Sepedomerus, and Sepedonea \ack a
basiphallus. The remaining genera have a
well-developed basiphallus.
12. Setal investment of foretibia: P (0)
without preapical setae; A (1) bearing a pair
of preapical setae.
The presence of a pair of preapical setae
on the foretibia is a synapomorphy for Oi-
dematops and Sciomyza.
13. Vestiture of the aedeagus (Figs. 214-—
217): P (O) without scalelike structures; A
(1) with scalelike structures.
In Calliscia and Parectinocera, the ae-
deagus is covered with small scalelike
structures, a synapomorphy for these gen-
era.
14. Proepisternal seta: P (0) present; A
(1) absent.
Steyskal (1965) used the presence of the
proepisternal seta to distinguish the tribe
Sciomyzini from the Tetanocerini. Most
genera of Tetanocerini lack this seta, al-
though Shannonia Malloch and Perilimnia
Becker possess one. For the latter genera
this condition is considered to be homo-
plastic.
15. Male terminalia (Figs. 57-59, 64-65,
68-160): P (O) with the 6th, 7th, and 8th
sternites separated, the 6th and 7th asym-
metrical; A (1) with the synsternite 7+8
and 6th sternite separated and asymmetri-
cal; A (2) with the synsternite 6+7+8 sym-
metrical.
According to Steyskal (1957) the asym-
metry of the male Acalyptratae postabdo-
men, including segments beyond the fifth,
is due to three basic movements within the
pupa: circumversion, reflection, and “‘stro-
phe.”’ The symmetry may occur as a result
of the obliteration of these movements.
Asymmetry in Sciomyzidae is restricted to
the sixth, seventh, and eighth segments.
There is a modification gradient of these
168
segments, changing from completely asym-
metrical to completely symmetrical, asso-
ciated to the fusion of the segments men-
tioned above.
16. Placement of sixth right abdominal
spiracle of the male (Figs. 68-114): P (0)
in membrane; A (1) in sclerotized tergite.
Sciomyzini and Eutrichomelina, Reno-
cera Hendel, and Ectinocera Zetterstedt
have the plesiomorphic condition of this
character, the spiracle is in the membrane.
From Antichaeta to Sepedonea, the spiracle
is in the tergite. In Ethiolimnia, there is a
reversal to the plesiomorphic condition.
17. Placement of seventh right abdominal
spiracle of the male (Figs. 68—114): P (O)
in membrane; A (1) in sclerotized tergite.
The plesiomorphic condition occurs in
most Sciomyzini except for Calliscia. From
Chasmacryptum Becker to Sepedonea most
genera have the spiracle in the tergite, ex-
cept for Dictyacium Steyskal and Ethiol-
imnia.
18. Shape of head: P (O) as in Fig. 52a;
A()-asin Figs s2bie’ @)as im Biesis2c;
A(8)-as inePiss 52d.
The head, from the ancestor to Chasma-
cryptum, has the shape shown in Fig. 52a.
Beyond the group of Shannonia + Perilim-
nia the intermediate apomorphic condition,
represented by Fig. 52c, is present. From
this state the other two apomorphic states
evolved. The pattern of head shape, as in
Fig. 52b, is present in Trypetolimnia and
Limnia Robineau-Desvoidy as homoplasies
and in Hedria, Dichetophora, Coremacera
Rondani, Dictyacium, and Euthycera La-
treille. The shape, as in Fig. 52d, defines
the group of Ethiolimnia, Thecomyia, Se-
pedoninus, Sepedonella, Sepedon, Sepe-
domerus, and Sepedonea. Because Teuto-
niomyia Hennig has a very different head,
compared to other genera, this character
was coded as missing in the matrix (Table
i):
19. Proportion between the length of the
pedicel and first flagellomere (Figs. 1—51):
P (QO) pedicel approximately half of first fla-
gellomere; A (1) pedicel subequal to the
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
length of the first flagellomere; A (2) ped-
icel approximately twice the length of the
first flagellomere.
The plesiomorphic condition is present
from Sciomyzini until Shannonia + Peri-
limnia. From this group to Ethiolimnia the
intermediate apomorphic condition (1) is
present. The apomorphic condition (2) is
present in the following genera: Teutoni-
omyia, Thecomyia, Sepedoninus, Sepedo-
nella, Sepedon, Sepedomerus, and Sepedo-
nea.
20. Condition of sternites of female ter-
minalia (Figs. 53-56): P (O) with sternites
6th, 7th, and 8th separate; A (1) with 6th
separate and 7th and 8th fused; A (2) 6th,
7th, and 8th incompletely fused; A (3) 6th,
7th, and 8th completely fused.
The transformation series was_ linear.
Two lineages arose from state 1. One of
these goes through the third state and the
second goes through the fourth state. The
evolution occurred from separated sternites
passing through steps of fusion to the com-
plete fusion of 6th, 7th, and 8th. All Scio-
myzini and the Tetanocerine genera until
Hoplodictya Cresson + Dictya Meigen
have sternites 6th, 7th, and 8th separate.
From the polytomy of Hydromya Robi-
neau-Desvoidy to the ancestor of Ethiol-
imnia the 7th and 8th sternites are fused.
Protodictya + Pherbecta and Poecilogra-
pha have the three sternites completely
fused as Sepedoninus, Sepedon, Sepedom-
erus, and Sepedonea. In Sepedonella, a ge-
nus belonging to this group, the three ster-
nites are separate. The monophyletic group
formed by Coremacera, Dictyacium, and
Euthycera, and the genera Limnia, Teuto-
niomyia, and Thecomyia have the sternites
6, 7, and 8 almost fused. Lines separating
these sternites are perceptible.
21. Aedeagus with lobed apex (Figs.
222-224, 230-238): P (O) absent; A (1)
present.
Ectinocera, Hoplodictya, and Perilimnia
+ Shannonia have the aedeagus with the
distal end lobed. This character apparently
appeared independently in these genera. In
VOLUME 113, NUMBER |
the other genera, the distal end of the ae-
deagus has several different shapes.
22. Subalar setae: P (0) absent; A (1)
present.
The subalar setae are present in Dictyo-
des, Ilione, Pherbina, Trypetoptera, Limnia,
Poecilographa, Pherbecta + Protodictya.
The apomorphic state apparently evolved
independently in Antichaeta.
23. Aedeagus convoluted as in Figs.
263-265, 281-286: P (0) absent; A (1)
present.
A convoluted aedeagus occurs in Proto-
dictya, Pherbecta, and Dictyodes. For Pro-
todictya and Pherbecta this character de-
fines the common ancestor.
24. Fourth abdominal sternite of the male
fused to fifth, forming a medioapical pro-
cess: P (OQ) absent; A (1) present.
The presence of a well-developed, me-
dioapical process in the fifth abdominal
sternite of the male as a complex structure
is a synapomorphy for the species of Pro-
todictya (Marinoni & Knutson 1992, Mar-
inoni & Carvalho 1993). In Pherbecta,
however, there is, at the same position, a
process that is less well developed and
which may be homologous to that of Pro-
todictya.
25. Lunula: P (0) not exposed; A (1) ex-
posed; A (2) greatly exposed.
In the outgroup, Pelidnoptera, genera of
Sciomyzini, and genera of Tetanocerini
from Eutrichomelina to Protodictya, the lu-
nula is not exposed. From Guatemalia to
Sepedonella, the intermediate state (1) is
present. In Ethiolimnia, there is a reversal
to the plesiomorphic state. In the apical
genera, Sepedon, Sepedomerus, and Sepe-
donea, the lunula is greatly exposed, which
is interpreted as an even more derived con-
dition from the intermediate state. In Teu-
toniomyia, which also has a greatly exposed
lunula, this character state is homoplastic.
26. Presutural supra-alar setae: P (0)
present; A (1) absent.
The plesiomorphic condition of this char-
acter is the presence of presutural supra-alar
setae. The absence of these setae is a syn-
169
apomorphy for the group Elgiva, Hedria,
Dichetophora, Ethiolimnia, Teutoniomyia,
Thecomyia, Sepedoninus, Sepedonella, Se-
pedon, Sepedomerus, and Sepedonea. In
Coremacera, Dictyacium, and Euthycera,
there is a reversal to the plesiomorphic
state. Tetanura has this seta, a condition
that is considered homoplastic.
27. Seventh and 8th abdominal tergites
of the female: P (0) separate; A (1) fused.
The fusion of the 7th and the 8th abdom-
inal tergites of the female is a character
state that links the genera Dichetophora,
Coremacera, Dictyacium, and Euthycera.
In Elgiva and Sepedonella, the fusion is ho-
moplastic.
28. Prominent eyes: (0) absent; (1) pres-
ent.
Ethiolimnia, Thecomyia, Sepedoninus,
Sepedonella, Sepedon, Sepedomerus, and
Sepedonea have prominent eyes, a syna-
pomorphy for these genera. As Ethiolimnia
is in a polytomy with Teutoniomyia, prom-
inent eyes are a confirmed synapomorphy
for Thecomyia, Sepedoninus, Sepedonella,
Sepedon, Sepedomerus, and Sepedonea and
only perhaps for Ethiolimnia.
29. Number of postalar setae: P (0) 2; A
(1) O-1.
The presence of two postalar setae is the
plesiomorphic condition. In Ectinocera and
in the genera beyond Ethiolimnia, the num-
ber of postalar setae is reduced to one or
none.
30. Ocellar setae: P (QO) present; A (1)
absent.
The presence of a pair of strong ocellar
setae has been considered a plesiomorphic
condition for the Muscomorpha (McAlpine
1989). In Pelidnoptera, these setae are pres-
ent, as they are in most Sciomyzidae, in-
cluding Salticella. These setae are absent in
Hedria and in the group of Thecomyia, Se-
pedoninus, Sepedonella, Sepedomerus +
Sepedonea.
31. Postpronotal setae: P (0) present; A
(1) absent.
Most Sciomyzidae have postpronotal se-
tae. The absence of these setae character-
170
izes the group of genera beyond Thecomyia.
In Dichetophora and Tetanura, the absence
of postpronotal setae is considered to be ho-
moplastic.
32. Number of ‘scutellar setae?“ PP")? 2
pairs; A (1) 1 pair.
Primitively the family Sciomyzidae has
two pairs of scutellar setae. One pair of scu-
tellar setae occurs in Dichetophora and the
group of genera beyond Thecomyia.
33. Number of notopleural setae: P (0) 2;
A (ty
Two notopleural setae are present in most
Sciomyzidae and in Pelidnoptera. Only the
genera Thecomyia, Sepedoninus, and Sepe-
donella have a single notopleural seta. A
reversal of this characters to the plesio-
morphic state of two notopleural setae oc-
curs in the genera Sepedon, Sepedomerus,
and Sepedonea.
34. Hindtibia arch-shaped: P (0) absent;
A (1) present.
The genera Teutoniomyia, Sepedoninus,
Sepedonella, Sepedon, Sepedomerus, and
Sepedonea have an arch-shaped hindtibia.
Only Thecomyia does not have the hindtibia
arch-shaped.
35. Head with sutures: P (0) distinct; A
(1) indistinct.
The head sutures of Thecomyia, Sepe-
doninus, Sepedonella, Sepedon, Sepedome-
rus, and Sepedonea are indistinct. This con-
dition is considered a synapomorphy for
this group.
36. Epandrium (Figs. 68—160): P (0) free;
A (1) fused to the 8th sternite.
The fusion of the epandrium to the 8th
sternite is a synapomorphy for the genera
Sepedonella, Sepedon, Sepedomerus, and
Sepedonea.
37. Hindtibia with a spinelike projection:
P (O) absent; A (1) present.
In most Sciomyzidae, there is no projec-
tion from the hindtibia. Sepedonea and Se-
pedomerus have a hindtibia that bears a
spinelike projection, which is a synapomor-
phy that links these two genera.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Discussion
Relationships.—The monophyly of the
Sciomyzidae is defined by a _ behavioral
character, the larvae are predators or para-
sitoids on aquatic or terrestrial Mollusca
(Berg 1953, Knutson et al. 1970, Griffiths
1972, Barnes 1979) and the presence of a
serrate ventral arch below the mouth hooks
in the larvae (Griffiths 1972). The serrate
ventral arch could be closely correlated
with the malacophagous feeding behavior.
The genus Salticella, representing the
Salticellinae, remains at the base of the
cladogram, being distinguished from the
subfamily Sciomyzinae and its monophyly
confirmed by the following three autapo-
morphies: four spermathecae; cell M nar-
rowed apically (this character was not used
in the analysis because of its variability
among genera of Sciomyzidae); and three
pairs of surstyli. Knutson et al. (1970),
working on the biosystematics of Salticella
fasciata (Meigen), concluded that this ge-
nus is more closely related to Sciomyzini
than to the Tetanocerini in morphological
and biological aspects. This relationship is
corroborated in the present analysis.
Among genera of the subfamily Scio-
myzinae, two main monophyletic clades
were formed, the first including Pherbellia,
Pteromicra, Calliscia, Parectinocera, Te-
tanura, Colobaea, Atrichomelina, Oidema-
tops, and Sciomyza; and the second includ-
ing the 40 remaining genera. The two
clades are concordant with the tribes Scio-
myzini and Tetanocerini of Steyskal (1965).
The genus Eutrichomelina, which tradition-
ally has been placed in the Sciomyzini, is
shown to be allied to the Tetanocerini in
this analysis.
Behavioral aspects of the larvae have
been discussed as possible synapomor-
phies for the tribes (Knutson & Lyneborg
1965, Abercrombie 1970). Toward one end
of the scale, where the Sciomyzini are
placed, the species that have been reared
are parasitoids on stranded aquatic or ter-
restrial mollusks (Colobaea (Knutson
VOLUME 113, NUMBER 1
0,5 mm
10
Figs. 1-11.
0,5 mm
2 ae es ?
7a
11
Right antenna, right lateral view: 1, Pelidnoptera; 2, Salticella; 3, Sciomyza; 4, Oidematops; 5,
Atrichomelina; 6, Tetanura;,; 7, Colobaea; 8, Calliscia; 9, Parectinocera, 10, Pteromicra; 11, Pherbellia. Fig.
11, scale = 1.0 mm; remaining Figs. scale = 0.5 mm.
1973); Pherbellia (Bratt et al. 1969); Pter-
omicra (Rozkosny & Knutson 1970); Scio-
myza (Foote 1959); Tetanura (Knutson
1970)). Toward the other end, most Tetan-
ocerini are predaceous on aquatic or ter-
restrial mollusks (Coremacera (Knutson
1973); Dichetophora (Vala et al. 1987);
Dictya (Valley & Berg 1977); Dictyodes
(Abercrombie 1970, Abercrombie & Berg
1978); Elgiva (Knutson & Berg 1964a,
Orth & Knutson 1987); Euthycera (Vala et
al. 1983, Vala & Caillet 1985); Hoplodic-
tya (Neff & Berg 1962); Hydromya (Knut-
son & Berg 1963); /lione (Knutson & Berg
1964b); Limnia (Vala & Knutson 1990);
Perilimnia (Kaczynski et al. 1969); Pher-
becta (Knutson 1972); Pherbina (Knutson
et al. 1975, Vala & Gasc 1990); Protodic-
tya (Abercrombie 1970, Neff & Berg
1961); Psacadina (Knutson et al. 1975);
Renocera (Foote & Knutson 1970); Sepe-
don (Neff & Berg 1966, Knutson et al.
172 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Ee
SM
eG a ai
12 13
0,5 mm
LEER OR
GA
24
0,5 mm
28
Figs. 12-29. Right antenna, right lateral view: 12, Eutrichomelina; 13, Ectinocera; 14, Renocera; 15, An-
tichaeta;,; 16, Chasmacryptum; 17, Shannonia; 18, Perilimnia; 19, Hoplodictya,; 20, Dictya; 21, Hydromya; 22,
Neolimnia; 23, Tetanoceroides; 24, Euthycerina; 25, Tetanocera; 26, Trypetolimnia; 27, Psacadina; 28, Steys-
kalina; 29, Dictyodes. Figs. 18 and 23 with scale 1.0 mm; the remaining Figs. 0.5 mm.
VOLUME 113, NUMBER 1 173
0,5 mm
1mm
1mm
wu G‘O
Figs. 30-43. Right antenna, right lateral view: 30, /lione; 31, Pherbina; 32, Trypetoptera; 33, Limnia; 34,
Poecilographa; 35, Pherbecta; 36, Protodictya; 37, Guatemalia; 38, Elgiva; 39, Hedria; 40, Dichetophora; 41,
Coremacera; 42, Dictyacium; 43, Euthycera. Figs. 30-35 and 40 with scale 0.5 mm; Figs. 37-39 and 41—43
with scale 1.0 mm.
1967); Sepedonea (Abercrombie 1970, 1970); Thecomyia (Abercrombie & Berg
Knutson & Valley 1978); Shannonia (Kac- 1975); Trypetoptera (Vala 1986)).
zynski et al. 1969); Tetanocera (Knutson A few species of Sciomyzini and Tetan-
et al. 1965); Tetanoceroides (Abercrombie ocerini, however, have an intermediate be-
174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs. 44-51.
Right antenna, right lateral view: 44, Ethiolimnia; 45, Teutoniomyia; 46, Thecomyia; 47, Se-
pedoninus,; 48, Sepedonella; 49, Sepedon; 50, Sepedomerus; 51, Sepedonea. Fig. 45 with scale 0.5 mm; the
remaining Figs. with scale 1.0 mm.
havior between these two behaviors, 1.e.,
sometimes they have parasitoid trends,
sometimes predaceous. Examples of these
kinds of behavior are evident in the species
Atrichomelina pubera Loew and Antichaeta
analis Zetterstedt. Atrichomelina pubera
belongs to the Sciomyzini. Depending on
circumstances this species has a predatory,
parasitoid or even saprophagous behavior
(Berg et al. 1959, Foote et al. 1960). Sap-
rophagy may be the primitive condition,
similar to that found in Salticella (Knutson
et al. 1970) and the ancestor of Sciomyzi-
dae (Abercrombie 1970). Antichaeta analis
has some parasitoid characteristics, such as
host specificity and oviposition on the host,
but its behavior is predatory with the larvae
feeding on snail eggs (Fisher & Orth 1964,
Knutson 1966, Knutson & Abercrombie
LOFT):
More studies related to the evolution of
the larval behavior are necessary to confirm
that the larval habit is responsible for re-
lationships among the genera within the
family.
Most genera in the analysis remained in
unresolved polytomies due to the great
number of homoplasies and low number of
synapomorphies.
Within the Sciomyzini, the relationships
of Pteromicra and Pherbellia are not well
resolved. RozkosSny & Knutson (1970),
studying the biology and immature stages
of Pteromicra, discovered that this genus is
closely related to Pherbellia. However,
even in our analysis, where these two gen-
era are in close proximity on the cladogram,
there is no adult character to corroborate
this relationship.
Knutson et al. (1990) considered Colo-
baea, Pherbellia, and Pteromicra to be re-
lated but did not elaborate any synapomor-
phies to support their grouping. In the same
paper they also suggest that Pherbellia is
paraphyletic.
Parectinocera and Calliscia appear in the
analysis as having the same ancestor. These
two genera are the only two within the
Sciomyzini with Neotropical distributions.
The genera Tetanura, Colobaea, Atricho-
VOLUME 113, NUMBER 1
52
0,5 mm
©»
Figs. 52—56.
syn. 6+7+8
Ce)
5
6
52. Shape of head: (a), State 0; (b), state 1; (c) state 2; (d) state 3. 53-56. Female terminalia,
ventral view: 53, Perilimnia; 54, Elgiva; 55, Coremacera; 56, Protodictya. Sternite 6 (st. 6); sternite 7 (st. 7);
sternite 8 (st. 8); synsternite 6+7+8 (syn. 6+7+8).
melina, Sciomyza, and Oidematops form a
clade established by characters 4 and 10
(Figs. 323-324). In this group Atrichome-
lina, Sciomyza + Oidematops have their re-
lationships well defined. The synapomorph-
ic character that links the three genera is the
presence of a well-developed male sub-
epandrial plate (Character 5). The other two
genera remain in polytomy.
In the Tetanocerini, Eutrichomelina, Ec-
tinocera, Renocera, Antichaeta, and Chas-
macryptum are at the base of the clade. Ec-
176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
1mm
bas.
( | 0,5 mm
62 63
Figs. 57-63. Male terminalia. Pelidnoptera. Aedeagal apodeme (aed. ap.); basiphallus (bas.); cerci (cer.);
aedeagal complex (aed. c.); epandrium (ep.); sternite 6 (st. 6); sternite 7 (st. 7); sternite 8 (st. 8); hypandrium
(hip.); paramere (par.); subepandrial plate (sub. pl.); anterior surstylus (a. s.); posterior surstylus (p. s.). Figs.
57—60 with scale 1.0 mm; Figs. 61—63 with scale 0.5 mm.
tinocera and Renocera remain in polytomy. already discussed by Fisher & Orth (1964)
Steyskal (1959), using intuitive analysis, & Knutson (1966). Adult characters suggest
considered Renocera, Chasmacryptum, and that this genus belongs to the Tetanocerini,
Tetanocera to be morphologically related. | while larval characters indicate its position
The uncertain position of Antichaeta was _ to be within the Sciomyzini. Boyes et al.
VOLUME 113, NUMBER 1 177
st. 8
/ Low cer.
ep.
oe
st. 7
= a.s
st. 6 a }
p.s
\ s.m
cs
p.s. 65
64
aed.
Figs. 64-67. Male terminalia. Salticella. Aedeagal apodeme (aed. ap.); cerci (cer.); aedeagal complex (aed.
c.); epandrium (ep.); sternite 6 (st. 6); sternite 7 (st. 7); sternite 8 (st. 8); hypandrium (hip.); paramere (par.);
anterior surstylus (a. s.); medial surstylus (s. m.); posterior surstylus (p. s.). Figs. 64—65 with scale 1.0 mm;
Figs. 66—67 with scale 0.5 mm.
(1969), studying the karyotype of Antichae-
ta melanosoma Melander, observed its sim-
ilarity with those of the Tetanocerini gen-
era. In the present analysis, Antichaeta is
placed in the Tetanocerini group of genera
near Chasmacryptum.
After Chasmacryptum, the clade of Shan-
nonia + Perilimnia appears as the sister
group of the other genera. Kaczynski et al.
(1969) studied the biosystematics of the im-
mature stages of these two genera and ob-
served morphological and behavioral affin-
ities between them. As observed by these
authors, the main similarities between the
larvae of Shannonia and Perilimnia are in
the cephalopharyngeal skeleton. They also
suggested the position of both genera with-
in the Tetanocerini.
178
Figs. 68-76.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Male terminalia, ventral view: 68, Sciomyza; 69, Oidematops; 70, Atrichomelina; 71, Tetanura;
72, Colobaea;, 73, Calliscia; 74, Parectinocera; 75, Pteromicra; 76, Pherbellia. Cerci (cer.); epandrium (ep.);
sternite 6 (st. 6); sternite 7 (st. 7); sternite 8 (st. 8); anterior surstylus (a. s.); posterior surstylus (p. s.); tergite 6
(terg. 6). Fig. 68 with scale 1.0 mm; the remaining Figs. with scale 0.5 mm.
Hoplodictya + Dictya form the next
clade. Boyes et al. (1969) compared its kar-
yotypes and recognized the proximity of
these two genera.
Hydromya, Neolimnia, and Tetanocero-
ides remain as a polytomy with three other
groups formed. The components of one of
these groups are: Euthycerina, Tetanocera,
Trypetolimnia, Psacadina, and Steyskalina.
The other group is formed by Dictyodes,
Ilione, Pherbina, Trypetoptera, Limnia,
Poecilographa, Pherbecta + Protodictya.
Within the latter group the genera Limnia,
Poecilographa, Pherbecta + Protodictya
are monophyletic. In the third group, which
comprises the last fifteen genera, we rec-
ognize two subgroups. One is formed by
Hedria, Dichetophora, Coremacera, Dic-
tyacium, and Euthycera, and the other by
Ethiolimnia, Teutoniomyia, Thecomyia, Se-
pedoninus, Sepedonella, and Sepedomerus
+ Sepedonea. The monophyly of the first
VOLUME 113, NUMBER I 179
0,5 mm
85
1mm
Figs. 77-87. Male terminalia, ventral view: 77, Ectinocera; 78, Renocera; 79, Antichaeta; 80, Chasmacryp-
tum; 81, Shannonia; 82, Perilimnia; 83, Hoplodictya; 84, Dictya; 85, Hydromya; 86, Neolimnia; 87, Euthycerina.
Cerci (cer.); epandrium (ep.); sternite 6 (st. 6); sternite 7 (st. 7); sternite 8 (st. 8); synsternite 7+8 (syn. 7+8);
synsternite 6+7+8 (syn. 6+7+8); anterior surstylus (a. s.); posterior surstylus (p. s.). Fig. 86 with scale 1.0 mm;
the remaining Figs. with scale 0.5 mm.
180 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
89 90
syn. 6+7+8
(So
QZG 93
0,5 mm
99
0,5 mm
Figs. 88-99. Male terminalia, ventral view: 88, Tetanocera; 89, Trypetolimnia; 90, Psacadina; 91, Steys-
kalina; 92, Dictyodes; 93, Ilione; 94, Pherbina; 95, Trypetoptera; 96, Limnia; 97, Poecilographa; 98, Pherbecta;
99, Protodictya. Cerci (cer.); epandrium (ep.); sternite 6 (st. 6); synsternite 7+8 (syn. 7+8); synsternite 6+7+8
(syn. 6+7+8); anterior surstylus (a. s.); posterior surstylus (p. s.). Figs. 90 and 98 with scale 0.5 mm; the
remaining Figs. with scale 1.0 mm.
VOLUME 113, NUMBER 1 181
0,5 mm 1mm
ep.
cer. Oy (s
103-7 ><
<a 102 ee
syn.7+8
110
109
pe 6+7+8 es
111 112 ain 113 AQ
0,5 mm
Figs. 100-114. Male terminalia, ventral view: 100-101, Guatemalia; 102, Elgiva; 103, Hedria; 104, Diche-
tophora; 105, Coremacera; 106, Dictyacium; 107, Euthycera; 108, Ethiolimnia; 109, Thecomyia; 110, Sepedon-
inus; 111, Sepedonella; 112, Sepedon; 113, Sepedomerus; 114, Sepedonea. Cerci (cer.); epandrium (ep.); sternite
6 (st. 6); synsternite 7+8 (syn. 7+8); synsternite 6+7+8 (syn. 6+7+8); anterior surstylus (a. s.); posterior
surstylus (p. s.). Figs. 100, 101, 104-106, 111—114 with scale 0.5 mm; the remaining Figs. with scale 1.0 mm.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
120
0,5 mm
123
cer.
Figs. 115-123. Male terminalia, left lateral view: 115, Sciomyza; 116, Oidematops; 117, Atrichomelina; 118,
Tetanura; 119, Colobaea; 120, Calliscia; 121, Parectinocera; 122, Pteromicra; 123, Pherbellia. Cerci (cer.);
epandrium (ep.); sternite 6 (st. 6); sternite 7 (st. 7); sternite 8 (st. 8); anterior surstylus (a. s.); posterior surstylus
(p. s.); tergite 6 (terg. 6). Fig. 115 with scale 1.0 mm; the remaining Figs. with scale 0.5 mm.
VOLUME 113, NUMBER 1 183
124
0,5 mm
Z 134 135
Figs. 124-135. Male terminalia, left lateral view: 124, Ectinocera; 125, Renocera; 126, Antichaeta; 127,
Chasmacryptum; 128, Shannonia; 129, Perilimnia; 130, Hoplodictya; 131, Dictya; 132, Hydromya; 133, Neo-
limnia; 134, Euthycerina; 135, Tetanocera. Cerci (cer.); epandrium (ep.); sternite 6 (st. 6); synsternite 7+8 (syn.
7+8); synsternite 6+7+8 (syn. 6+7+8); anterior surstylus (a. s.); posterior surstylus (p. s.). Figs. 133 with scale
1.0 mm; the remaining Figs. with scale 0.5 mm.
184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs. 136-146. Male terminalia, left lateral view: 136, Trypetolimnia; 137, Psacadina; 138, Steyskalina; 139,
Dictyodes; 140, Ilione; 141, Pherbina; 142, Trypetoptera; 143, Limnia; 144, Poecilographa; 145, Pherbecta;
146, Protodictya. Cerci (cer.); epandrium (ep.); sternite 6 (st. 6); synsternite 7+8 (syn. 7+8); synsternite 6+7+8
(syn. 6+7+8); anterior surstylus (a. s.); posterior surstylus (p. s.). Figs. 138 and 146 with scale 0.5 mm; the
remaining Figs. with scale 1.0 mm.
VOLUME 113, NUMBER |
0,5 mm
syn. 6+7+8 7
mm
i ase 2
syn. 6+7+8
<a ae
Figs. 147-160. Male terminalia, left lateral view: 147, Guatemalia; 148, Elgiva; 149, Hedria; 150, Diche-
tophora; 151, Coremacera; 152, Dictyacium; 153, Euthycera; 154, Ethiolimnia; 155, Thecomyia; 156, Sepedon-
inus; 157, Sepedonella; 158, Sepedon; 159, Sepedomerus; 160, Sepedonea. Cerci (cer.); epandrium (ep.); sternite
6 (st. 6); synsternite 7+8 (syn. 7+8); synsternite 6+7+8 (syn. 6+7+8); posterior surstylus (p. s.). Figs. 148—
150, 154, 155 with scale 1.0 mm; the remaining Figs. 0.5 mm.
186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs. 161-163. Male terminalia, posterior view: 161, Sciomyza; 162, Oidematops; 163, Atrichomelina. Cerci
(cer.); subepandrial plate (sub. pl.); anterior surstylus (a. s.); posterior surstylus (p. s.).
subgroup, from Thecomyia to Sepedonea, is group comprising the genera Thecomyia,
established by eight characters: 6, 19, 28, Sepedon, Sepedonella, Sepedoninus, Sepe-
30, 31, 32, 33 and 35. It is the most cor- domerus, and Sepedonea. Steyskal (1973:
roborated lineage within this analysis. 143) stated that “I do not believe that this
Steyskal (1973) placed Sepedon in a_ group is sufficiently distinct from more typ-
VOLUME 113, NUMBER 1
0,5 mm
164
187
Figs. 164-169.
Male terminalia, posterior view: 164, Tetanura; 165, Colobaea; 166, Calliscia; 167, Parec-
tinocera; 168, Pteromicra; 169, Pherbellia. Cerci (cer.); subepandrial plate (sub. pl.); anterior surstylus (a. s.);
posterior surstylus (p. s.).
ical Tetanocerini, especially from such gen-
era as Hedria and Dichetophora, to be giv-
en the rank of tribe or even subtribe. We
would prefer to call it merely “‘the Sepedon
group.” ”’ This statement agrees with the
result of the present analysis, but with the
addition the genera Ethiolimnia and Teu-
toniomyia to the Sepedon group.
Proposal of classification.—The follow-
ing natural classification was done using the
sequencing convention (Wiley 1981). The
subfamilies Salticellinae and Sciomyzinae
and the tribes Sciomyzini and Tetanocerini
are confirmed.
Sciomyzidae Fallén, 1820
Salticellinae Hendel, 1924
Salticella Robineau-Desvoidy, 1830
Sciomyzinae Fallén, 1820
Sciomyzini Fallén, 1820
Pherbellia Robineau-Desvoidy, 1830
188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
p.s.
| 171
, 173
= :
175
174 176
Figs. 170-176. Male terminalia, posterior view: 170, Antichaeta; 171, Shannonia; 172, Perilimnia; 173, Hoplod-
ictya; 174, Dictya (lateral view), 175, Hydromya; 176, Neolimnia. Anterior surstylus (a. s.); posterior surstylus (p. s.).
0,5 mm
a.S.
Pteromicra Lioy, 1864 Eutrichomelina Steyskal, 1975
Calliscia Steyskal, 1975 Ectinocera Zetterstedt, 1838
Parectinocera Becker, 1919 Renocera Hendel, 1900
Colobaea Zetterstedt, 1837 Antichaeta Haliday, 1838
Tetanura Fallén, 1820 Chasmacryptum Becker, 1907
Atrichomelina Cresson, 1920 Shannonia Malloch, 1933
Oidematops Cresson, 1920 Perilimnia Becker, 1919
Sciomyza Fallén, 1820 Hoplodictya Cresson, 1920
Tetanocerini Hendel, 1900 Dictya Meigen, 1803
VOLUME 113, NUMBER 1 189
177
0,5 mm
” 181
182
Figs. 177-186. Male terminalia, posterior view: 177, Euthycerina; 178, Tetanocera; 179, Trypetolimnia;
180, Psacadina; 181, Steyskalina; 182, Dictyodes; 183, Ilione; 184, Pherbina; 185, Trypetoptera; 186, Limnia.
Anterior surstylus (a. s.); posterior surstylus (p. s.).
190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
189
ip [
a.s. Nace |
‘ic
(‘ 7
p.s
191
190
. 7 mn
192 193
194
: =—ias
p.s.
195 Wee
Figs. 187-196. Male terminalia, posterior view: 187, Poecilographa; 188, Pherbecta; 189, Protodictya; 190,
Guatemalia; 191, Elgiva; 192, Hedria; 193, Dichetophora; 194, Coremacera; 195, Dictyacium; 196, Euthycera.
Anterior surstylus (a. s.); posterior surstylus (p. s.).
Hydromya Robineau-Desvoidy, 1830 Tetanocera Duméril, 1800
Neolimnia Barnes, 1976 Trypetolimnia Mayer, 1953
Tetanoceroides Malloch, 1933 Psacadina Enderlein, 1939
Euthycerina Malloch, 1933 Dictyodes Malloch, 1933
VOLUME 113, NUMBER 1 19]
198
Ke ‘
Figs. 197-203. Male terminalia, posterior view: 197, Ethiolimnia; 198, Thecomyia; 199, Sepedoninus, 200, Se-
pedonella; 201, Sepedon; 202, Sepedomerus; 203, Sepedonea. Anterior surstylus (a. s.); posterior surstylus (p. s.).
Ilione Verbeke, 1964 Ethiolimnia Verbeke, 1950
Pherbina Robineau-Desvoidy, 1830 Teutoniomyia Hennig, 1952
Trypetoptera Hendel, 1900 Thecomyia Perty, 1833
Limnia Robineau-Desvoidy, 1830 Sepedoninus Verbeke, 1950
Poecilographa Melander, 1913 Sepedonella Verbeke, 1950
Pherbecta Steyskal, 1956 Sepedon Latreille, 1804
Protodictya Malloch, 1933 Sepedomerus Steyskal, 1973
Guatemalia Steyskal, 1960 Sepedonea Steyskal, 1973
Elgiva Meigen, 1838 Steyskalina Knutson, 1999
Hedria Steyskal, 1954
Dichetophora Rondani, 1868
Coremacera Rondani, 1856
Dictyacium Steyskal, 1920 The subfamilies Salticellinae and Scio-
Euthycera Latreille, 1829 myzinae are monophyletic. In the Sciomy-
Conclusions
192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
205 207
209 210
«
208
212
213
Figs. 204-213. Male terminalia. 204, 208, 206, 210, 212, Right lateral view; 205, 207, 209, 211, 213, frontal
view. 204, 205, Sciomyza; 206, 207, Oidematops; 208-209, Atrichomelina; 210-211, Tetanura; 212—213, Co-
lobaea. Aedeagal apodeme (aed. ap.); basiphallus (bas.); aedeagus (aed.); gonopod (gon.); hypandrium (hip.);
paramere (par.).
VOLUME 113, NUMBER 1
>
>
.
.
“e
ve
vee
eve
7 We |
216
218
219
221
Male terminalia. 214, 216, 218, 220, Right lateral view; 215, 217, 219, 221, frontal view.
Figs. 214-221.
214, 215, Calliscia; 216, 217, Parectinocera; 218, 219, Pteromicra; 220, 221, Pherbellia. Aedeagal apodeme
(aed. ap.); basiphallus (bas.); aedeagus (aed.); gonopod (gon.); hypandrium (hip.); paramere (par.).
194 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
par.
aed.
0,5 mm
227 228
: 229
aed.
>,
< 33
ee Youd vis
¥
"weve vec
Othe? 1 2
ae 3 1
230
Figs. 222—232. Male terminalia. 222, 225, 227, 230, Right lateral view; 223, 226, 228, 231, frontal view;
224, 229, 232, left lateral view. 222-224, Ectinocera; 225—226, Antichaeta; 227—229, Chasmacryptum; 230-
232, Shannonia. Aedeagal apodeme (aed. ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.).
VOLUME 113, NUMBER 1
aed.
aed.
0,5 mm
Figs. 233-241.
195
234
241
Male terminalia. 233, 236, 239, Right lateral view; 234, 237, 240, frontal view; 235, 238,
241, left lateral view. 233-235, Perilimnia; 236—238, Hoplodictya,; 239-241, Dictya. Aedeagal apodeme (aed.
ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.).
zinae the tribes Sciomyzini and Tetanocer-
ini are monophyletic.
The tribe Sciomyzini includes Pherbel-
lia, Pteromicra, Calliscia, Parectinocera,
Colobaea, Tetanura, Atrichomelina, Oide-
matops, and Sciomyza.
The tribe Tetanocerini includes Antichae-
ta, Chasmacryptum, Coremacera, Dicheto-
Phora, Dictya, Dictyacium, Dictyodes, Ec-
tinocera, Elgiva, Ethiolimnia, Euthycera,
Euthycerina, Eutrichomelina, Guatemalia,
Hedria, Hydromya, Hoplodictya, Ilione,
196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
0,5 mm
NO
BSS
NO
aed.ap.
< =e
247
245 246
249 250
253
252
Figs. 242-253. Male terminalia. 242, 245, 248, 251, Right lateral view; 243, 246, 249, 252, frontal view;
244, 247, 250, 253, left lateral view. 242-244, Hydromya; 245-247, Neolimnia; 248-250, Euthycerina; 251-—
253, Tetanocera. Aedeagal apodeme (aed. ap.); aedeagus (aed.); paramere (par.).
VOLUME 113, NUMBER 1 197
0,5 mm
par.
257 258 259
es
261 ES) Fe
aed.
265
263
Figs. 254-265. Male terminalia. 254, 257, 260, 263, Right lateral view; 255, 258, 261, 264, frontal view;
256, 259, 262, 265, left lateral view. 254—256, Trypetolimnia; 257-259, Psacadina; 260—262, Steyskalina; 263-—
265, Dictyodes. Aedeagal apodeme (aed. ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.).
198 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ew
273 274
Figs. 266-274. Male terminalia. 266, 269, 272, Right lateral view; 267, 270, 273, frontal view; 268, 271,
274, left lateral view. 266—268, Ilione; 269-271, Pherbina; 272-274, Trypetoptera. Aedeagal apodeme (aed.
ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.).
Limnia, Neolimnia, Perilimnia, Pherbecta, nea, Shannonia, Steyskalina, Tetanocera,
Pherbina, Poecilographa, Protodictya, Tetanoceroides, Trypetoptera, Trypetolim-
Psacadina, Renocera, Sepedoninus, Sepe- nia, Teutoniomyia, and Thecomyia.
donella, Sepedon, Sepedomerus, Sepedo- Within the Sciomyzini a monophyletic
VOLUME 113, NUMBER | 199
SP
279
par.
284 286
Figs. 275—286. Male terminalia. 275, 278, 281, 284, Right lateral view; 276, 279, 282, 285, frontal view;
377, 380, 383, 386, left lateral view. 275-277, Limnia; 278-280, Poecilographa; 281-283, Pherbecta; 284-286,
Protodictya. Aedeagal apodeme (aed. ap.); aedeagus (aed.); paramere (par.).
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
aed.ap.
aed.
£
E
287 288 3 | 4
par.
200
298
y aed.ap.
bas. OJ
eR 29 , 295
Figs. 287-298. Male terminalia. 287, 290, 293, 296, Right lateral view; 288, 291, 294, 297, frontal view
289, 292, 295, 298, left lateral view. 287-289, Guatemalia; 290-292, Elgiva; 293-295, Hedria; 296-298, Di-
chetophora. Aedeagal apodeme (aed. ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.)
VOLUME 113, NUMBER 1
0,5 mm
aed.
201
301
300
Figs. 299-307. Male terminalia. 299, 302, 305, Right lateral view; 300, 303, 305, frontal view; 301, 304,
306, left lateral view. 299-301, Coremacera; 302-304, Dictyacium; 305-307, Euthycera. Aedeagal apodeme
(aed. ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.).
group was formed by the genera Colobaea,
Tetanura, Atrichomelina, Oidematops +
Sciomyza. In this group there is the sub-
group formed by Atrichomelina, Oidema-
tops + Sciomyza.
The groups of Shannonia + Perilimnia
and Hoplodictya + Dictya, at the base of
Tetanocerini, are monophyletic. Three
other major groups of genera are formed
within the Tetanocerini: the first by Eu-
202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
>
>
Dias
SOS
2B
>
ie
?
2r5?
>?
0,5 mm
312 313
Figs. 308-315. Male terminalia. 308, 311, 314, Right lateral view; 309, 312, 315, frontal view; 310, 313,
left lateral view. 308-310, Ethiolimnia; 311-313, Thecomyia; 314, 315, Sepedoninus. Aedeagal apodeme (aed.
ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.).
thycerina, Psacadina Enderlein (1933), dictya, and the third by Guatemalia, El-
Steyskalina, Tetanocera, and Trypetolim- giva, Dichetophora, Hedria, Coremacera,
nia, the second by the genera Dictyodes,- Dictyacium, Euthycera, Ethiolimnia, Teu-
Ilione, Pherbina, Trypetoptera, Limnia, toniomyia, Thecomyia, Sepedoninus, Se-
Poecilographa, and Pherbecta + Proto- pedonella, Sepedon, Sepedomerus + Se-
VOLUME 113, NUMBER 1 203
316 317
0,5 mm
—— aed.ap.
ee par.
aed.
‘318
322
Figs. 316-322. Male terminalia. 316, 318, 320, 322, Right lateral view; 317, 319, 321, frontal view. 316,
317, Sepedonella; 318, 319, Sepedon; 320, 321, Sepedomerus; 322, Sepedonea. Aedeagal apodeme (aed. ap.);
aedeagus (aed.); paramere (par.).
pedonea. Within the last group the genera _ ships; and the group of Ethiolimnia, Teu-
Coremacera, Dictyacium, and Euthycera toniomyia, Thecomyia, Sepedoninus, Se-
form a monophyletic clade without any pedonella, Sepedon, Sepedomerus + Se-
resolution of their phylogenetic relation- pedonea is monophyletic.
nN
C= ie)
nN a Nn
— nN =
= =
oe
= ez
_
©-2 On Rw rm)
= Mm-0 2 = =~
—
—
ey
—
—_
ra)
g
_
~
_
_
GC}
XN =a =
BA me YES
©
_
_
~ At
=)
_
o
nN
—
—
=
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SS
-_
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a
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o =
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—_
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RROD Ses el
O©ONSOO& N
- SOON |] W&
=- Nn N
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=- = = -—- = _
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= N
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oud
on;,DM WY
w
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_
Fig 2323:
bpe '€z
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Pelidnoptera PA
Salticella PA AT
Sciomyza NA PA
Oidematops NA
Atrichomelina NA NT
Tetanura PA
Colobaea NA PA AT
Calliscia NT
Parectinocera NT
Pteromicra NA PA
Pherbellia NA PA NT AT OR AS
Eutrichomelina NT
Ectinocera PA
Renocera NA PA
Antichaeta NA PA
Chasmacryptum PA
Shannonia NT
Perilimnia NT
. Hoplodictya NA NT
Dictya NA PA NT
Hydromya PA
Neolimnia AS
Tetanoceroides NT
Euthycerina NT
Tetanocera NA PA OR
Trypetolimnia PA
Psacadina PA
Steyskalina OR
Dictyodes NT
llione PA NA OR
Pherbina PA
Trypetoptera NA PA
Limnia NA PA
Poecilographa NA
Pherbecta NA
Protodictya NT
Guatemalia NT
Elgiva NA PA
Hedria NA
Dichetophora PA OR AS
Coremacera PA
Dictyacium NA
Euthycera NA PA NT
Ethiolimnia AT
Teutoniomyia NT
Thecomyia NT
Sepedoninus AT
Sepedonella AT
Sepedon NA PA NT AT OR AS
Sepedomerus NT OR
Sepedonea NT
Consensus cladogram. Outgroup Pelidnoptera. 37 characters. Consistency index: 0.60. Retention
index: 0.88. Regions: AT, Afrotropical; AS, Australian; NA, Nearctic; NT, Neotropical; OR, Oriental; PA, Pa-
learctic.
VOLUME 113, NUMBER 1 205
OL
db}
BL
6L
Pelidnoptera
1} {| tt | | | Satticelta
| | | Sciomyza
Atrichomelina
tT } | t—4+--4 }
| | |
| 4h a feel Tetanura
}
Colobaea
|_|
| | | | | Oidematops
were IT TT] 111) camiseta
| | | Parectinocera
tied Seles | | Pteromicra
Pherbellia
Eutrichomelina
g | | Ectinocera
| | | Renocera
| | Antichaeta
ey Chasmacryptum
| Shannonia
hele Perilimnia
Hoplodictya
Pee Dictya
| | Hydromya
‘z
Neolimnia
Tetanoceroides
Euthycerina
Tetanocera
Trypetolimnia
Psacadina
Steyskalina
| | | Dictyodes
| llione
Pherbina
Trypetoptera
|
Limnia
| | Poecilographa
Pherbecta
Protodictya
Guatemalia
ROMER RT BASE me! = = Ta Re ae ae es as a
Elgiva
Hedria
Dichetophora
-
|
|
+
=
Coremacera
1
|
Dictyacium
Euthycera
Ethiolimnia
|
Teutoniomyia
1
i
Thecomyia
Sepedoninus
Sepedonelia
Sepedon
=|a
SEIMESESaIeE
Sepedomerus
Sepedonea
Fig. 324. Arrangement of character states among genera of Sciomyzidae. White = plesiomorphic condition;
gray (light and dark) = intermediate and relatively more apomorphic conditions for multistate characters (pro-
gressing from light gray to dark gray); black = apomorphic condition.
Acknowledgments
Many individuals and institutions have
contributed to this analysis. We thank the
following institutions and respective cura-
tors for their cooperation in lending speci-
mens.
BMNH _ The Natural History Museum,
London, England
Colecao de Entomologia Pe. J.
S. Moure, Department of Zool-
ogy, Universidade Federal do
Parana, Curitiba, Parana, Brazil
Instituto Nacional de Pesquisas
da Amazonia, Manaus, Ama-
zonas, Brazil
Museu Nacional, Universidade
Federal do Rio de Janeiro, Rio
de Janeiro, Rio de Janeiro, Bra-
zil
Museu de Zoologia da Univer-
sidade de Sao Paulo, Sao Paulo,
Sao Paulo, Brazil
University of California, River-
side, California, U.S.A.
former United States National
Museum, collections in the Na-
tional Museum of Natural His-
tory, Smithsonian Institution,
Washington, D.C., U.S.A.
We thank Dr. Claudio J. B. de Carvalho,
Dr* Marcia S. Couri, Dr* Dilma S. Napp, &
Dr. Angelo P. do Prado for reviewing the
manuscript and Dr. John Kingsolver for re-
viewing the English version. We also thank
Drs. Rudolf RozkosSny & Lloyd V. Knutson
for very constructive discussions on nu-
merous aspects and the conclusions that are
presented in this paper.
DZUP
INPA
MNRJ
MZSP
UCR
USNM
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Appendix 1.
List of genera and type species used in
the analysis.
Antichaeta analis Meigen, 1830
Atrichomelina pubera (Loew), 1862
Calliscia calliscelles (Steyskal), 1963
Chasmacryptum seriatimpunctatum Becker,
1907
Colobaea bifasciella (Fallén), 1820
Coremacera marginata (Fabricius), 1775
Dichetophora obliterata (Fabricius), 1805
Dictya umbrarum (Linnaeus), 1758
Dictyacium ambiguum (Loew), 1864
Dictyodes dictyodes (Wiedemann), 1830
Ectinocera borealis Zetterstedt, 1838
Elgiva cucularia (Linnaeus), 1767
Ethiolimnia platalea Verbeke, 1950
Euthycera chaerophylli (Fabricius), 1798
Euthycerina vittithorax Malloch, 1933
Eutrichomelina fulvipennis (Walker), 1837
Guatemalia straminata (Wulp), 1897
Hedria mixta Steyskal, 1954
Hoplodictya setosa (Coquillett), 1901
Hydromya dorsalis (Fabricius), 1775
209
Ilione albiseta (Scopoli), 1763
Limnia unguicornis (Scopoli), 1763
Neolimnia repo Barnes, 1976
Oidematops ferrugineus Cresson, 1920
Parectinocera neotropica Becker, 1919
Perilimnia albifacies Becker, 1919
Pherbecta limenitis Steyskal, 1956
Pherbellia punctata (Fabricius), 1791
Pherbina coryleti (Scopoli), 1763
Poecilographa decora (Loew), 1864
Protodictya chilensis Malloch, 1933
Psacadina disjecta Enderlein, 1939
Pteromicra glabricula (Fallén), 1820
Renocera pallida (Fallén), 1820
Salticella fasciata (Meigen), 1830
Sciomyza simplex Fallén, 1820
Sepedomerus macropus (Walker), 1849
Sepedon sphegea (Fabricius), 1775
Sepedonea lindneri (Hendel), 1932
Sepedonella nana Verbeke, 1950
Sepedoninus planifrons Verbeke, 1950
Shannonia meridionalis Zuska, 1969
Steyskalina picta Ghorpadé and Marinoni,
1999
Tetanocera elata (Fabricius), 1781
Tetanoceroides mesopleuralis Malloch,
1933
Tetanura pallidiventris Fallén, 1820
Teutoniomyia plaumanni Hennig, 1952
Thecomyia longicornis Perty, 1833
Trypetolimnia rossica Mayer, 1953
Trypetoptera punctulata (Scopoli), 1763
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):210—217. 2000.
Review of the chewing louse genus Abrocomophaga
(Phthiraptera: Amblycera), with description of two new species
Roger D. Price and Robert M. Timm
(RDP) Department of Entomology and Plant Pathology, Oklahoma State University,
Stillwater, Oklahoma 74078-0464, U.S.A. (Current address) 4202 Stanard Circle,
Fort Smith, Arkansas 72903-1906, U.S.A.;
(RMT) Natural History Museum and Department of Ecology & Evolutionary Biology,
University of Kansas, Lawrence, Kansas 66045-2454, U.S.A.
Abstract.—The South American chewing louse Abrocomophaga chilensis
Emerson & Price has, since its description, remained the sole known member
of the amblyceran family Abrocomophagidae. We herein provide a redescrip-
tion of the species as well as descriptions and illustrations for two new species
of Abrocomophaga: A. emmonsae off Cuscomys ashaninka Emmons from Pert
and A. hellenthali off the degu, Octodon degus (Molina), from Chile. After our
reevaluation of the status of the family Abrocomophagidae, we consider it a
synonym of the family Gyropidae.
Resumen.—Desde su descripcion original, el piojo sudamericano Abrocom-
ophaga chilensis Emerson & Price, ha permanecido como el inico miembro
conocido de la familia Abrocomophagidae, perteneciente a los ambliceros. En
este trabajo presentamos una redescripcion de dicha especie y ademas descrip-
ciones e ilustraciones de dos especies nuevas del género Abrocomophaga: A.
emmonsae parasito de Cuscomys ashaninka Emmons proveniente del Pert y
A. hellenthali parasito del degu, Octodon degus (Molina), de Chile. Después
de reevaluar el status de la familia Abrocomophagidae, la consideramos como
sin6nimo de la familia Gyropidae.
Emerson & Price (1976) described a dis-
tinctive new taxon of South American
chewing louse, Abrocomophaga chilensis,
and placed it in a new monotypic family,
the Abrocomophagidae. All known speci-
mens of this louse were found on a single
chinchilla rat, Abrocoma bennetti Water-
house, from Chile. Since the original de-
scription, little has been written about this
enigmatic and phylogenetically important
amblyceran (Emerson & Price 1985), al-
most certainly because no new information
was available. We recently obtained series
of Abrocomophaga from two other South
American rodent hosts; these lice represent
two new species in this genus. With the ad-
dition of these two new species, we reeval-
uate the status of the family Abrocomo-
phagidae and the genus Abrocomophaga.
Our purpose here is to describe and illus-
trate these new species and to establish the
synonymy for the family Abrocomophagi-
dae.
Family Gyropidae Kellogg
Gyropidae Kellogg, 1896:68. Type genus:
Gyropus Nitzsch.
Abrocomophagidae Emerson & Price,
1976:425. Type genus: Abrocomophaga
Emerson & Price. New synonymy.
The features associated with the species
of Gyropidae and the relationship of this
family to other members of the suborder
Amblycera have been accurately treated by
Clay (1970) and will not be repeated here.
VOLUME 113, NUMBER 1
In her treatment of the amblyceran Phthir-
aptera, Clay (1970) followed the organiza-
tion presented by Ewing (1924) in recog-
nizing the Gyropidae as being divided into
three subfamilies: (1) the Gyropinae with
six pairs of abdominal spiracles (on III-—
VIII) and at least one pair of legs having a
modified tarsal claw strongly adapted for
clasping hair; (2) the Gliricolinae with only
five pairs of abdominal spiracles (on II-—
VII) and legs with a single greatly reduced
tarsal claw; and (3) the Protogyropinae with
six pairs of abdominal spiracles (on II-—
VIII) and all legs having a single unmodi-
fied tarsal claw.
The establishment of the family Abro-
comophagidae by Emerson & Price (1976)
was based primarily on the fact that Abro-
comophaga chilensis, the sole member of
this new family, had an unmodified tarsal
claw on each leg and only five pairs of ab-
dominal spiracles, a combination of char-
acters seen in no other gyropid louse. How-
ever, as our understanding of character evo-
lution has matured over the years, we have
come to the conclusion that these differenc-
es are of generic-level rather than familial-
level significance. In fact, the main differ-
ence between the Abrocomophagidae and
the monotypic Protogyropinae was that the
former had only five pairs of abdominal
spiracles (on III—VII), whereas the latter
had six pairs (on III—VIII). Because the
number of abdominal spiracles in the Gy-
ropidae varies from the primitive six pairs
to a reduced state of five pairs, it is our
feeling now that this difference is not of
familial significance and that the Abrocom-
ophagidae should be considered a synonym
of the family Gyropidae (subfamily Proto-
gyropinae) as delineated by Clay (1970). A
discussion of other features associated with
the Abrocomophagidae and Abrocomopha-
ga may be found in Emerson & Price
(1976). Lakshminarayana (1976) presented
a thorough review of the suprageneric clas-
sification of the Phthiraptera and gave a list-
ing of the superfamily, family, subfamily,
and tribe names proposed to that time.
211
Genus Abrocomophaga Emerson & Price
Abrocomophaga Emerson & Price, 1976:
425. Type species: Abrocomophaga chi-
lensis Emerson & Price.
This genus is separated from others in the
family Gyropidae (and suborder Ambly-
cera) in having known representatives with
all legs having an unmodified tarsal claw
and the abdomen with only five pairs of spi-
racles.
Abrocomophaga chilensis Emerson &
Piice
Abrocomophaga chilensis Emerson &
Price, 1976:426. Type host: Abrocoma
bennetti bennetti Waterhouse.
Male.—Emerson & Price (1976) provide
a full illustration in Fig. 5 and two aspects
of the male genitalia in Figs. 3 and 4. Head
with numerous medium-length setae on
dorsal and ventral surfaces; without heavy
dorsal seta near antennal base. Pronotum
with 10—14 setae; prosternum with 5-7 se-
tae, mesosternum with 8—10. Metanotum
with 10 setae, including 6 longer median
setae and each side laterally with 1 long, 1
short seta; metasternal plate with 12—14 se-
tae. Abdominal tergal setae: I, 4; II, 6—8;
III, 8—10; IV, 10-12; V, 11-13; VI, 11-14;
VII, 10-12; VIII, 6-9. Terga III—VII each
with small faint median pigmentation; V—
VIII each with median pair of shorter setae
recessed posterior to main row. Posterior
margin of each of pleura IIJ—VIII with 1
very long and 1 adjacent short heavy seta.
Last tergum with 2 short, 1 very long seta
on each side. Abdominal sternal setae: II,
7-9; Il, 9-12; IV, 11-15; V, 12-14; VI,
10—13; VII, 9—11. Sterna IV—VII each with
small faint median pigmentation. Subgeni-
tal plate with 7 or 8 medium to short me-
dian setae. Genitalia with only ill-defined
weak mesosomal sclerites associated with
sac and with blunt apical tip of parameres.
Dimensions (in mm): temple width, 0.19—
0.20; head length, 0.16—0.18; prothorax
width, 0.15—0.16; metathorax width, 0.18—
212
0.20; abdomen width at segment V, 0.30—
0.31; genitalia width, 0.08; genitalia length,
0.17—0.20; genitalia paramere length, 0.06—
0.07; total length, 0.94—0.98.
Female.—Emerson & Price (1976) pro-
vide a full illustration in Fig. 2 and one of
the ventral terminalia in Fig. 1. Head and
thorax as for male, abdomen differing as
follows. Tergal setae: IV, 9-11; V, 10-12;
VI, 10-13; VII, 9-12. All setae on terga V—
VIII aligned in row. Last tergum with 2
very long setae on each side, in addition to
2 shorter setae. Sternal setae: II, 8—10; III,
10-13; IV, 11-13; V, 11-14; VI, 11-13;
VII, 8-11. Subgenital plate with 9-11 me-
dioanterior setae and convex posterior mar-
gin bearing median fringe of small spicules.
Anal fringe ventrally of 5 + 5 and dorsally
of 3 + 3 longer lateral setae, with few mi-
nute median setae in each row. Dimensions
(an mm): temple width, 0.20—0.22; head
length, 0.17—0.20; prothorax width, 0.16—
0.17; metathorax width, 0.19—0.20; abdo-
men width at segment V, 0.32—0.35; anus
width, 0.07—0.08; total length, 1.07—1.14.
Material.—10 male, 10 female paratypes
of Abrocomophaga chilensis, ex Abrocoma
bennetti bennetti, Chile.
Remarks.—This species is recognized by
both sexes with consistently small dimen-
sions, including narrow head and abdomen,
and their abdominal tergal and sternal setal
counts; the female with a narrow anus; and
the male with unique genitalia and a small
recessed pair of abdominal tergal setae on
each of V-—VIII. Type deposited in the
NMNH.
Emerson & Price described Abrocomo-
phaga chilensis on the basis of a large se-
ries of specimens from a single host indi-
vidual of Abrocoma bennetti bennetti from
Chile collected by Robert E. Martin. We
can now provide more specific information
on the type host and type locality. The type
host collected by Robert E. Martin on 2 No-
vember 1974 bears his field number 1105
and is deposited in the Recent mammal col-
lection at the Field Museum in Chicago as
FMNH 119794 with locality information as
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
follows: Chile: Santiago Province; 10 km
W of Til Til [Tiltil; 1000 m]. The host was
an adult female.
Abrocomophaga emmonsae, new species
Figs. 1—4
Type host.—Cuscomys ashaninka Em-
mons.
Male.—As in Fig. 1. Differing from
Abrocomophaga chilensis as follows. Ab-
dominal tergal setae: IV, 14; V, 18; VI, 14;
VI, 16. Terga I-VII each with small me-
dian faintly pigmented area; V—VII each
with median pair of shorter setae recessed
posterior to main row. Abdominal sternal
setae: IT, 10; III, 14; IV, 16; V, 15; VI-VII,
12. Sterna I—VII each with lightly pig-
mented median area. Genitalia (Fig. 3) with
prominent inwardly curved sharply pointed
parameres and conspicuous associated me-
sosomal sclerites as shown. Dimensions (in
mm): temple width, 0:26; head length, 0.20;
prothorax width, 0.19; metathorax width,
0.26; abdomen width at segment V, 0.40;
genitalia width, 0.09; genitalia length, 0.20;
genitalia paramere length, 0.06; total
length, 0.98.
Female.—As in Fig. 2. Differing from
Abrocomophaga chilensis as follows. Ab-
dominal tergal setae: IV, 12—13; V, 12-16;
VI, 13-18; VII, 13-16. Abdominal sternal
setae: II, 8-11; III, 11-15; IV, 13-17; V,
13—16; VI, 12—18; VII, 11-14. Ventral ter-
minalia as in Fig. 4. Dimensions (in mm):
temple width, 0.26—0.27; head length,
0.20—0.22; prothorax width, 0.19—0.20;
metathorax width, 0.23—0.26; abdomen
width at segment V, 0.41—0.47; anus width,
0.09—0.10; total length, 1.06—1.16.
Type material.—Holotype male, ex Cus-
comys ashaninka, Peré: Cuzco, Cordillera
Vilcabamba: (11°39'36'S5/73°38'3 1 WwW) sek
3350 m, 15 June 1997, coll. Louise H) Em-
mons #1359; in collection of the Museo de
Historia Natural, Universidad Nacional de
San Marcos, Lima, Peri, MUSM 12715 °.
Paratypes: 6 females, same data as holo-
type; in collections of the Universidad Na-
VOLUME 113, NUMBER 1
213
Lae
Figs. 1-4. Abrocomophaga emmonsae: (1) Dorsal-ventral male; (2) Dorsal-ventral female; (3) Male geni-
talia; (4) Ventral female terminalia.
cional de San Marcos and the National Mu-
seum of Natural History, Washington, D.C.
Etymology.—This new species is named
in honor of Louise H. Emmons, National
Museum of Natural History, Smithsonian
Institution, who collected the host and en-
thusiastically encouraged the junior author
to examine it for ectoparasites. Her survey
efforts in South America have greatly con-
tributed to our understanding of this fauna
and her field guides to Neotropical mam-
mals have created a broader understanding
214
of the rainforest for both the public and stu-
dents of all ages, and have certainly con-
tributed to conservation efforts.
Remarks.—Although this new species is
close to Abrocomophaga chilensis, both
sexes are readily separable by their consis-
tently larger dimensions, including broader
head and abdomen, and their larger number
of abdominal tergal and sternal setae; the
female with the broader anus; and the male
with the recessed pair of setae on only ab-
dominal terga V—VII and with unique gen-
italia that have the distinctive mesosomal
sclerites and acutely pointed parameres.
Abrocomophaga hellenthali, new species
Figs. 5-8
Type host.—Octodon degus (Molina).
Male.—As for Abrocomophaga chilensis
except as follows. Head (Fig. 6) with heavy
dorsal seta near antennal base. Abdomen
(Fig. 5) with tergal setae: II, 5—6; III, 6—7;
IV-—V, 7-8; VI-VII, 8; VUI, 5—6. Terga I-
VII each with small faint median pigmen-
tation; V and VII each with median pair of
shorter setae recessed posterior to main
row. Posterior margin of each of pleura II-—
VIII with short slender seta adjacent to very
long seta. Last tergum with | long, 3-4
short setae on each side. Abdominal sternal
setae: II, 6; II, 8—9; IV, 9-10; V, 8-10; VI,
7-8; VII, 6. Sterna II-VII each with small
faint median pigmentation. Genitalia (Fig.
7) with numerous conspicuous spinelike
mesosomal sclerites associated with sac and
with sharply pointed apical tip of parameres.
Dimensions (in mm): temple width,
0.23—0.26; head length, 0.18—0.20; protho-
rax width, 0O.17—0.18; metathorax width,
0.22—0.23; abdomen width at segment V,
0.38—0.41; genitalia width, 0.08—0.09; gen-
italia length, 0.19—0.21; genitalia paramere
length, 0.07; total length, 0.83—0.88.
Female.—Head and thorax as for male,
abdomen (Fig. 8) differing as follows. Ter-
gal setae: VIII, 6—7. All setae on terga V
and VII aligned in row. Last tergum with 2
very long setae on each side, in addition to
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
2 shorter setae. Sternal setae: II, 6—8; VI,
8-9; VII, 6—7. Subgenital plate with 9-11
medioanterior setae and convex posterior
margin bearing median fringe of small spic-
ules. Anal fringe as for Abrocomophaga
chilensis. Dimensions (in mm): temple
width, 0.24—0.26; head length, 0.19—0.20;
prothorax width, 0.17—0.19; metathorax
width, 0.22—0.26; abdomen width at seg-
ment V, 0.43—0.47; anus width, 0.09—0.10;
total length, 0.94—0.98.
Type material.—Holotype male, ex Oc-
todon degus, Chile: Santiago, Santiago, 2.5
km NE of Cerro Manquehue, Trappist Mon-
astery, 9 July 1975, coll. Robert E. Martin
#1222 (FMNH 119614 &); in collection of
the K. C. Emerson Entomology Museum,
Oklahoma State University, Stillwater.
Paratypes: 2 males, 3 females, same data as
holotype; 1 female, same except 1 August
1975, Robert E. Martin #1235 (FMNH
119756 3); 1 male, 1 female, same except
27 April 1976, Robert E. Martin #1432
(FMNH 119639 <); all in same collection
as holotype.
Etymology.—This species is named for
Ronald A. Hellenthal, University of Notre
Dame, in recognition of his strong interest
in chewing louse taxonomy and his many
years of cooperative studies with RDP.
Remarks.—This species is separated
from the other two of the genus by both
sexes having large dimensions, including a
broad head and abdomen, a heavy dorsal
preantennal head spine, a short slender seta
adjacent to the very long seta on each of
pleura IHI—VIII, and a consistently small
number of abdominal tergal and sternal se-
tae; the female with a broad anus; and the
male with a recessed pair of setae only on
abdominal terga V and VII and with unique
genitalia that have a distinctive assemblage
of spinous mesosomal sclerites and acutely
pointed parameres.
Discussion
The caviomorph rodent family Octodon-
tidae contains 6 genera and 11 living spe-
VOLUME 113, NUMBER 1
6 ae SS ees SE \
et
thats ele sey
Li ee eee ry
Sian |
“76
Figs. 5-8. Abrocomophaga hellenthali: (5) Dorsal-ventral male abdomen; (6) Dorsal male head; (7) Male
genitalia; (8) Dorsal-ventral female abdomen.
cies, all with very restricted Andean or pre-
Andean ranges; Octodon degus is the most
widely distributed species. With the discov-
ery of Cuscomys ashaninka, the cavio-
morph family Abrocomidae now contains
two Recent genera and a total of five spe-
cies (Emmons 1999). Both families occupy
a diverse array of high-elevation habitats,
and their geographic ranges overlap broadly
in southern South America. The relation-
ships between the Abrocomidae and Octo-
dontidae have been the subject of debate.
The abrocomids were historically treated as
a subfamily of the Echimyidae and more
recently as either a subfamily of the Octo-
dontidae or as a closely related family in
the superfamily Octodontoidea following
Reig (1986) and Carleton (1984). However,
Glanz & Anderson (1990) suggested that
the abrocomids are more closely aligned
with the chinchillas and may belong in the
superfamily Chinchilloidea.
Where accurate records are available, we
have found gyropid lice to be extremely
host-specific ectoparasites (Price & Timm
1997), with speciation of lice closely par-
alleling speciation in their mammalian
hosts. The various species in a louse genus
are almost always restricted to the various
species of a host genus or to very closely
related genera of hosts. Our discovery of
the two new species of Abrocomophaga de-
scribed herein, although clearly supporting
the host specificity we observe in gyropids,
is odd in that closely related species of lice
are found on different families of rodents.
Two of the species of Abrocomophaga now
known—A. chilensis and A. emmonsae—
are apparently host-specific parasites of
abrocomid rodents (Abrocoma bennetti and
Cuscomys ashaninka, respectively) and one
species—A. hellenthali—is apparently a
host-specific parasite of an octodontid ro-
dent (Octodon degus). Because only three
species are known in the genus Abrocom-
ophaga and the genus is only known from
five separate host individuals in three sep-
216
arate genera, it is premature to attempt to
reconstruct a phylogeny for the group.
Naturally occurring transfers (host
switching) of parasitic lice between nonre-
lated hosts is not common, but has been
documented in both mammals and birds
(Paterson et al. 1999). Nest sharing between
different species of mammals is rare in na-
ture; however, sequential use of tree hole
nests by birds is widespread. Timm (1983)
postulated that the broad host distributions
of species and genera seen in many lineages
of bird lice are in part due to host transfers
that might occurred during the evolutionary
history of the groups. Mammal lice exhibit
greater host specificity because opportuni-
ties for lice to colonize new host taxa are
rare. Although nest sharing by different
species of rodents is exceedingly rare, it has
been well documented for Octodon degus
and Abrocoma bennetti in Chile. In a field
study with marked individuals, Fulk (1976)
found O. degus and A. bennetti regularly to
share burrows and even the same nests. In
excavating nests, he found female A. ben-
netti with their own infants as well as infant
degus. Correspondingly, female O. degus
also had both species of young in their
nests. Fulk (1976:504) considered nest
sharing a common behavior in these two
species, and suggested “‘This practice may
be mutualistic, each animal contributing to
the survival of the other’s young.”
Although we don’t know that nest shar-
ing occurs between Octodon degus and
Abrocoma bennetti throughout their range,
the fact that it does occur demonstrates how
an ancestral Abrocomophaga could be
transferred from one host family to another.
With only three species of Abrocomophaga
known to date, we cannot differentiate be-
tween the hypotheses that the current dis-
tribution of species we see on the rodent
families Abrocomidae and Octodontidae
are historically shared or relatively recent
host transfers. When additional species of
Abrocomophaga from other abrocomid and
octodontid hosts are known, we may be
able to address whether the Abrocomidae or
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
the Octodontidae, or perhaps a common an-
cestor, represent the ancestral host for this
enigmatic and poorly known group of lice.
Acknowledgments
We thank Louise Emmons and Linda
Gordon, National Museum of Natural His-
tory, Washington, D.C., for allowing us to
remove specimens of lice from mammal
specimens housed at the National Museum,
thereby making the discovery of Abrocom-
ophaga emmonsae and other new species
possible. Larry Heaney and Bruce Patterson
made our work at the Field Museum pos-
sible and productive. Robert Anderson ex-
amined specimens of Abrocoma for us as
well as translated our Abstract into Spanish
for the Resumen included herein. Rob An-
derson, Barbara Clauson, Louise Emmons,
Marcela Gomez-Laverde, and Ricardo Pal-
ma provided valuable assistance on earlier
drafts of this manuscript. This manuscript
has been approved for publication by the
Director, Oklahoma Agricultural Experi-
ment Station, Stillwater.
Literature Cited
Carleton, M. D. 1984. Introduction to rodents. Pp.
255-265 in S. Anderson and J. K. Jones, Jr.,
eds., Orders and families of Recent mammals
of the world. John Wiley & Sons, New York,
686 pp.
Clay, T. 1970. The Amblycera (Phthiraptera: Insec-
ta).—Bulletin of the British Museum (Natural
History), Entomology 25:73-—98.
Emerson, K. C., & R. D. Price. 1976. Abrocomophag-
idae (Mallophaga: Amblycera), a new family
from Chile.—The Florida Entomologist 59:
425-428.
. 1985. Evolution of Mallophaga on mammals.
Pp. 233-255 in K. C. Kim, ed., Coevolution of
parasitic arthropods and mammals. John Wiley
& Sons, New York, 800 pp.
Emmons, L. H. 1999. A new genus and species of
abrocomid rodent from Peru (Rodentia: Abro-
comidae).—American Museum Novitates 3279:
1-14.
Ewing, H. E. 1924. On the taxonomy, biology, and
distribution of the biting lice of the family Gy-
ropidae.—Proceedings of the United States Na-
tional Museum 63:1—42.
Fulk, G. W. 1976. Notes on the activity, reproduction,
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and social behavior of Octodon degus.—Journal
of Mammalogy 57:495—505.
Glanz, W. E., & S. Anderson. 1990. Notes on Bolivian
mammals. 7. A new species of Abrocoma (Ro-
dentia) and relationships of the Abrocomidae.—
American Museum Novitates 2991:1—32.
Kellogg, V. L. 1896. New Mallophaga, I, with special
reference to a collection made from maritime
birds of the Bay of Monterey, California.—Pro-
ceedings of the California Academy of Sciences
(Series 2) 6:31—168.
Lakshminarayana, K. V. 1976. Nomenclatural changes
in Phthiraptera—some suggestions.—Ange-
wandte Parasitologie 17:160—167.
Paterson, A. M., R. L. Palma, & R. D. Gray. 1999.
How frequently do avian lice miss the boat?
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Implications for coevolutionary studies.—Sys-
tematic Biology 48:214—223.
Price, R. D., & R. M. Timm. 1997. A new subgenus
and four new species of Gliricola (Phthiraptera:
Gyropidae) from the Caribbean hutias (Roden-
tia: Capromyidae).—Proceedings of the Biolog-
ical Society of Washington 110:285-—300.
Reig, O. A. 1986. Diversity patterns and differentiation
of high Andean rodents. Pp. 404—440 in F. Vuil-
leumier and M. Monasterio, eds., High altitude
tropical biogeography. Oxford University Press,
New York, 649 pp.
Timm, R. M. 1983. Fahrenholz’s rule and resource
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Pp. 225-265 in M. H. Nitecki, ed., Coevolution.
University of Chicago Press, Chicago, 392 pp.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):218—223. 2000.
A new species of the genus Cubacubana
(Insecta: Zygentoma: Nicoletiidae)
from a Mexican cave
Luis Espinasa
CEAMISH, Universidad Autonoma del Estado de Morelos, Av. Universidad 1001 Col. Chamilpa,
Cuernavaca, Morelos CP 62210, Mexico
Abstract.—A new cavernicolous species of the genus Cubacubana (Zygen-
toma: Nicoletiidae) is described. This species is probably more distant from
any other species of this genus described until now.
The genus Cubacubana was originally
described with three cavernicolous species
from Cuba (Wygodzinsky & Hollinger
1977), although Espinasa (1999b) has pro-
posed that one of the species is of the genus
Anelpistina (Silvestri, 1905). In Wygodzin-
sky’s paper, he asserted that the Mexican
fauna of free-living and cavernicolous Ni-
coletiines is very rich and quite possibly
species of Cubacubana will also be found
on the mainland. Later another Cubacubana
species was described from Aruba island
(Mendes 1986) and finally a species was
described from México, C. mexicana (Es-
pinasa, 1991), thus fulfilling Wygodzin-
sky’s prediction. Further exploration of
caves in México has shown that organisms
of the genus Cubacubana are actually abun-
dant (Espinasa 1989).
The new species has a series of charac-
ters that permit us to define it as a cave
adapted troglobite such as the overall large
size, enlarged appendages, and surface
crawling as opposed to occurring under
rocks. Its presence in at least two different
cave localities can be explained because
they are within the same karstic area and
probably are a single underground system
(Diamant-Adler 1991).
Materials and Methods
Detailed descriptions of the caves can be
found in the “‘Tepeyollotli’’ bulletins of the
SMES speleological society (Diamant-Ad-
ler 1991). Organisms collected were crawl-
ing on the cave’s floor. They were placed
into a vial with 96% ethanol. Dissections
were made with the aid of a stereo micro-
scope and the different parts of the body
were mounted in fixed preparations with
Hoyer’s solution. All illustrations were
made with aid of a camera lucida attached
to a compound microscope.
Types were deposited in the following
collection: LESM-DB-MEX (Laboratory of
Ecology and Systematic of Microarthro-
pods. Department of Biology, Faculty of
Sciences, UNAM. México D.E). Catalog
number: ZYG-5.
Cubacubana asymmetrica, new species
Figs. 1[A—G, 2A—F
Type material.—México, Puebla State,
Coyomeapan municipality, Tepepa, ““TP4-
13” Cave (780 meters deep and more than
5 kilometers long). Ex soil, 23 Dec 1990,
L. Espinasa col. Male holotype, two male
paratypes and five female paratypes.
Other localities: México, Puebla State,
Coyomeapan municipality, Huitzilatl,
‘““Xaltégoxtl’”> Cave (60 meters deep and
more than 2 kilometers long). Ex soil, 3 Jan
1991, R. Espinasa col. Two males and two
females.
Description.—Maximum body length
22.5 mm. Maximum length of antennae 34
VOLUME 113, NUMBER 1
mm, of caudal appendages 35 mm. When
complete, antennae measure 3X length of
body and caudal appendages 2 length of
body. General color light yellow to white.
Macrochaetae simple or forked.
Head with approximately 5 + 5 macro-
chaetae on border of insertion of antennae.
Antennae of male with 83 preserved arti-
cles; distal articles very slender and long,
with internal sensory organs similar to C.
negreai (Wygodzinsky & Hollinger, 1977).
Basal article of antennae in males without
projections. Pedicellus of antennae of male
elongate as shown in Fig. 1B, with clusters
of unicellular glands arranged in 5 groups.
In adults, right pedicellus bigger than left
and longer than basal article (Fig. 1B—C).
Female basal articles of antennae simple.
Mouthparts very elongate (Figs. 1A and
2A); galea almost as long as lacinia (Fig.
2A), with 2 apical conules of different
width (Fig. 2B). Two teeth on lacinia and
on bigger tooth’s base, an extra very small
hyaline tooth. Labial palp long and slender,
apical article one and a half times longer
than wide and subtriangular (Fig. 1A). La-
bium and Ist article of labial palp with
macrochaetae. Mandibles with 4 macro-
chaetae, without small pegs on the bigger
tooth. Legs elongate; hind tibia approxi-
mately 8—9 times longer than wide. Leg
chaetotaxy as in C. mexicana (Espinasa,
1991). Claws of normal size.
Cerci of male with a very short basal ar-
ticle, a very long 2nd one bearing numerous
spines, followed by numerous short articles
of simple chaetotaxy. In adults, the very
long article is slightly curved and with
spines, all inserted on tubercles and roughly
of the same size and form (Fig. 1D—E). The
spine row does not reach base of article.
Female cerci simple.
Thorax with short macrochaetae: 2 + 2
on anterior border of pronotum, 3 + 3 mac-
rochaetae on nota’s lateral borders, and | +
1 submedian distinct macrochaetae apart
from several setae of varied sizes on nota’s
posterior borders.
Urotergite X of both sexes with a small
219
degree of prominence and posterior part
short in length, approximately one fifth its
width, with a shallow emargination (Fig.
1G), posterior angles with a few macro-
chaetae of varied sizes, length of inner ma-
crochaetae almost equal to distance be-
tween them.
Abdominal terga and sterna as in other
members of genus. Abdominal sterna H—
VII subdivided into coxites and sternite.
Sterna VIII and [X of male entire. Uroster-
num III and IV of male without modified
coxites. Urosternum VIII of male shallowly
emarginate on posterior margin, angles of
emargination pointed to slightly rounded.
Urosternum IX of male straight behind,
without modifications (Fig. 1F). Stylets H-—
VIII with two macrochaetae and an extra
subapical pair. Stylets [X larger than others,
with four macrochaetae and the extra sub-
apical pair. Terminal spine with small teeth.
In males and females styles IX without
spines.
Penis and parameres as shown in Fig. 1E
Parameres very short, broaden slightly at
base on inner face, and attaining only % of
stylets IX. Surface of parameres with short
setae, apical portion with numerous very
short setae. Subgenital plate of female sub-
elliptic or parabolic slightly rounded, slight-
ly longer than wide. Ovipositor surpassing
apices of stylets IX by a distance equal to
% the length of stylets (Fig. 2F). Gonapo-
physes with approximately 17 articles.
Post-embryonic development as in Ta-
bles 1—2, Figures 1D—E and 2C—EF Length
of body can be obtained from the length of
hind tibia according to the next formula:
Length of body = (5.57) length of hind
Gbiaate 65 3
Etymology.—asymmetrica = Asymmet-
ric. Makes reference to the asymmetric size
of pedicellus in males, the right one being
longer than the left.
Remarks.—Cubacubana asymmetrica
can be differentiated from other members
of the subfamily Cubacubaninae by the fol-
lowing characters: Species of genus Texo-
Nw
N
o
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
eS
t. set
L | \ Pay | Lee
Wore ell | =
va if == 4 f |! ot = z
ea |
bi fs “is peste) x: eae
! A ! ote 24 -
Fl bi aX
1
! Y
wotant
!
ue
2,
i}
1
1
'
!
'
Ot
cin
i]
at
i]
L at ae
DiI IPN es
yan ne
v Ase vx
aE ection Ne
Moris “i ic
ae iN si
ant Bid Viti
! WAY
uw TaN ES
rig j+\
14 iW
ml a ary
ii) D aS
1
x
S42
Fig. 1. Cubacubana asymmetrica n. Sp.: A, labium with palp; B, Male. Basal portion of left antenna; C,
Male. Basal portion of right antenna; D, Juvenile male. Basal portion of cercus with five spines; E, Adult male.
Basal portion of cercus with seven spines; K Male. Genital area; G, Urotergum X.
reddellia (Wygodzinsky, 1973) or Squami-_ icoletia (Paclt, 1979) has a rugged endo-
gera (Espinasa, 1999a), have scales, which podium, absent in Cubacubana. Prostheci-
are lacking in Cubacubana. Allonicoletia na (Silvestri, 1933), has a submentum with
(Mendes, 1992) lacks stylets in urosternite conspicuous lateral lobes bearing numerous
II, which are present in Cubacubana. Neon- glandular pores, absent in Cubacubana.
VOLUME 113, NUMBER I
Xk |
ee |
Pint.
[onennnann-==]
B
ames
5
ee
O
Fig) 2:
221
|
-1 mm.
[cet tr ere
Cubacubana asymmetrica n. Sp.: A, Maxilla; B, Apical portion of maxilla; C-K Post-embryonic
development of female. Ovipositor and subgenital plate. Length of ovipositor in Table 2 as follows: C = —%,
DS sal, E. = 0,.Fi— hh.
Most species of Anelpistina, have articulat-
ed submedian appendages in urosternite IV
of males, which are absent in Cubacubana.
From the two species of Anelpistina that
lack such appendages; A. decui (Wygodzin-
sky & Hollinger, 1977) and A. inappendi-
cata (Espinasa, 1999b), and from the de-
scribed species of Cubacubana; C. ramosi
(Wygodzinsky, 1959), C. negreai (Wygod-
zinsky & Hollinger, 1977), C. arubana
(Mendes, 1986), and C. mexicana (Espina-
sa, 1991), C. asymmetrica differs by the
posterior part of urotergite X, which is short
in length (one fifth the width of the uroter-
gite), while in the other species it is longer
(approximately one half to one third).
Males are also differentiated by long pedi-
cellus (twice or more long than wide) and
also by the asymmetry of adults, in which
right pedicellus is bigger than left.
Cubacubana asymmetrica can be further
differentiated from C. mexicana, the only
N
N
Table 1.—Post-embryonic development of male Cubacubana asymmetrica.
MALES:
(Fig. 1D-E)
Spines in stylets
IX
Spines in Cerci
Parameres vs.
stylets IX
Sensory cones
urosternum IX
Emargination
urosternum VIII
Spines in III
urosternum
Length both
pedicellus
Length
Hind
tibia
3/4 No
1/2
No 1/3
Yes
Yes
Unequal No NOES No Small
Equal
mm
Cave of -“iR4-137"
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
+++ +4
++ +4 +4
1.50
2.74
Sa10
Sn8
oo
Cave of ‘*Xaltégoxtl’’
2.74
2.88
a
Table 2.—Post-embryonic development of female
Cubacubana asymmetrica.
FEMALES:
Ovipositor surpassing apex stylets IX
: (Fig. 2C—F)
Lenethvhind. ‘= Se ees
tibia mm = Sy? =i (0) 1/4
Caver or, “TP4213~
1.74 45
3.45 ae
3.45 ae
3.49 il
Cave of *‘Xaltégoxtl’’
1.76 ==
2.89 =
3.16 at
other Mexican Cubacubana species de-
scribed, by its longer antenna (three times
the length of the body instead of one and a
half), caudal appendages (two times the
length of the body instead of slightly more
than one), maxillary palp (distal article 11
or more times longer than wide instead of
less than ten) and legs (hind tibia approxi-
mately eight to nine times longer than wide
instead of six). Furthermore males do not
have spines or cones on urosternum III and
IX or on styles IX. Gonapophyses of fe-
males have fewer articles (approximately
17 instead of 22).
Cubacubana asymmetrica shares char-
acteristics present in other genera, such as
the curvature of cerci of Prosthecina ad-
dititia (Wygodzinsky, 1951), or some type
of asymmetry in the pedicellus as in Cole-
tinia subterranea (Silvestri, 1902) and Col-
etinia asymetrica (Bach de Roca et al.,
1985). These characters are probably the re-
sult of convergent evolution.
Acknowledgments
I thank Dr. José G. Palacios-Vargas, di-
rector of “‘Laboratorio de Ecologia y siste-
matica de Microartr6podos”’, where most of
the descriptive work was done and the di-
rector of CEAMISH-UAEM,, Dr. Oscar Do-
rado, for support this research. Thanks are
also due to Dr. Luis EK Mendes and Monika
Baker for reviewing the manuscript.
VOLUME 113, NUMBER 1
Literature Cited
Bach de Roca, C., L. E Mendes, & M. Gaju Ricart.
1985. Sur une nouvelle espéce et une nouvelle
citation de Nicoletiinae de Sierra Morena (Cor-
doue, Espagne).—Bolletino della Societa ento-
mologica italiana 117(8—10):132—140.
Diamant-Adler, R. 1991. Participacién de la S.M.E.S.
en MEXPE IV, expedicion de la S.Q.S. a la Si-
erra Negra.—Tepeyollotli: Gaceta de la Socie-
dad Mexicana de Exploraciones Subterraneas 5:
22-28.
Espinasa, L. 1989. Bioespeleologia.—Tepeyolloth:
Gaceta de la Sociedad Mexicana de Explora-
ciones Subterraneas 4:45—46.
. 1991. Descripci6n de una nueva especie del
género Cubacubana (ZYGENTOMA: Nicoleti-
idae) y registro del género para América Con-
tinental.—Folia Entomolégica Mexicana 82:5—
16.
. 1999a. A new genus of the subfamily Cuba-
cubaninae (Insecta: Zygentoma: Nicoletiidae)
from a Mexican cave.—Proceedings of the Bi-
ological Society of Washington 112:52-58.
. 1999b. Two new species of the genus Anel-
pistina (Insecta: Zygentoma: Nicoletiidae) from
Mexican caves, with redescription of the ge-
nus.—Proceedings of the Biological Society of
Washington 112(1):59-69.
Mendes, L. FE 1986. Nouvelles données sur le Zygen-
toma (Insecta) de 1’ Amerique centrale et du Me-
xique.—Bulletin du Muséum national d’ Histoire
naturelle,. Paris (4) 8 (A) (2):333-—342.
. 1992. Novos dados sobre os tisanuros (Micro-
Ze5
coryphia e Zygentoma) da América do Norte.—
Garcia de Orta 16(1—2):171-193.
Paclt, J. 1979. Neue Beitrage zur Kenntnis der Apter-
ygoten-Sammlung des Zoologischen Instituts
und Zoologischen Museums der Universitat
Hamburg. VI Weitere Doppel- und Borstensch-
wanze (Diplura: Campodeidae. Thysanura: Lep-
ismatidae und Nicoletiidae).—Entomologische
Mitteilungen aus dem zoologischen Museum
Hamburg 6(105):221-—228.
Silvestri, EK 1902. Materiali per lo studio dei Tisanuri.
III. Nuove specie di Nicoletia.—Bolletino della
Societa entomologica italiana 33:223-—227.
. 1905. Materiali per lo studio dei Tisanuri. VI.
Tre nuovo sottogenero.—Redia (Firenze) 2:
111-120.
. 1933. Nuovo contributo alla conoscenza dei
tisanuri de Mesico.—Bolletino del Laboratorio
di Zoologia general e agraria di Portici 27:127—
144.
Wygodzinsky, P. 1951. Apuntes sobre “‘Thysanura”’
americanas.—Acta zoologica Lilloana 11:435-—
458.
. 1959. Contribution to the knowledge of the
“Thysanura” and ‘“‘Machilidae’’ (Insecta).—
Revista Brasileira de Biologia 19(4):441—457.
. 1973. Description of a new genus of cave
Thysanura from Texas (Nicoletiidae. Thysanu-
ra. Insecta).—American Museum Novitates
2518:1-8.
, & A. M. Hollinger. 1977. A Study of Nico-
letiidae from Cuba (Thysanura).—Resultats des
Expéditions Bioespeleologiques Cubano-rou-
maines 4 Cuba 2:313-324.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):224—237. 2000.
Three new species of bathyal cidaroids (Echinodermata: Echinoidea)
from the Antarctic region
Rich Mooi, Bruno David, F Julian Fell, and Thérése Choné
(RM) Department of Invertebrate Zoology & Geology, California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118-4599;
(BD & TC) UMR CNRS 5561, Université de Bourgogne, 6, bd. Gabriel, F-21000, Dijon, France;
(FJF) Box 222, Errington, British Columbia, Canada VOR 1V0O
Abstract.—Two new species of Aporocidaris, A. eltaniana and A. usarpi,
and one new species of Notocidaris, N. lanceolata, are described from material
collected during United States Antarctic Research Program expeditions in the
1960’s. All three species occur in the bathyal zone near or south of the Sub-
antarctic Convergence. Gonopore sizes and peristomial morphology suggest
that these species are sexually dimorphic and that the females can brood direct-
developing young. However, only one of the Aporocidaris species was found
to harbor embryos on the peristomial membrane.
In the mid- to late 1960’s, Dr. Richard H.
Chesher, at the Museum of Comparative
Zoology at Harvard (MCZ), studied echi-
noid material collected under the auspices
of the United States Antarctic Research
Program (USARP). The USARP, which be-
gan in 1955, included trawling and dredg-
ing expeditions by ships in the vicinity of
Antarctica (Fig. 1). Material collected by
the U.S. National Science Foundation Re-
search Vessel R/V Eltanin is of particular
importance to this paper. Dr. Chesher iden-
tified numerous USARP echinoids, in the
process discovering several new species. In
some cases, he proposed manuscript names
and indicated type specimens for new spe-
cies (for example, see David & Mooi 1990),
but he was unable to continue systematic
work after 1967, thereby leaving these
names and manuscripts unpublished.
Chesher’s cidaroid research was incorporat-
ed into a dissertation (Fell 1976), and two
of the species described below were cov-
ered in this work. As a result, several pu-
tative types for which no names have been
published are deposited in the National Mu-
seum of Natural History (NMNH)-the ma-
jor repository of echinoid collecting during
the USARP, and in the MCZ. Here, we pub-
lish these names for the first time.
Antarctic cidaroids are represented by
approximately 20 species in 5 genera, and
all belong to the subfamily Ctenocidarinae.
With one or two exceptions, the ctenoci-
darines are restricted to the Antarctic and
Subantarctic regions and are among the
most diverse of the Antarctic echinoids. Al-
though their taxonomy was hugely ad-
vanced in Fell (1976), an exact count of
valid taxa is difficult, given the uncertain
specific and subspecific status applied to the
variants. We are currently engaged in a da-
tabasing project that will result in a com-
prehensive overview of all Antarctic echi-
noids, including cidaroids, and it was this
review that prompted the work on the taxa
described herein.
Much of the Antarctic cidaroid fauna still
requires revision in a modern context. For
example, the only recent attempt (Smith &
Wright 1988) to resolve phylogenetic rela-
tionships among any of the cidaroids con-
centrated on Cretaceous forms, and did not
deal with Antarctic taxa. Phylogenetic re-
lationships among the different genera and
species are still speculative and will remain
VOLUME 113, NUMBER 1
- 4
90°
New Zealand 0
180° (0
Fip- 1.
TOT
225
@ Aporocidaris eltaniana
A Aporocidaris usarpi
¥% Notocidaris lanceolata
7 NUMBER OF
DEPTH STATIONS
, (m) » ea SAR ae |
\ (0)
2:
a L
:
\
Sr
1000 @
90°
\
\
\
2000
vt
S
xv
4000
Collecting localities and recorded depths (graph at right) for Aporocidaris eltaniana new species, A.
usarpi new species, and Notocidaris lanceolata new species. Number of stations refers to the number of R/V
Eltanin stations that for a given species fall within the depth range indicated by the height of the shaded box.
so until a more complete knowledge of the
taxonomy is established. The lack of con-
sistent, well-delineated features as well as
detailed analyses of morphological varia-
tion makes the Antarctic cidaroids a partic-
ularly difficult group, as evidenced by the
almost unusable keys provided by previous
major revisions such as Mortensen (1928).
In addition, material is often rare, and sev-
eral taxa are represented only by juveniles,
or have not been collected since they were
first described from single or very few in-
dividuals. Therefore, determination of ter-
minal taxa to be used in phylogenetic anal-
ysis is in its infancy, and any effort to place
the taxa described here in an evolutionary
context is premature.
Jackson (1912) and Smith (1984) provid-
ed excellent comparative overviews of ci-
daroid morphology. Cidaroids are very easy
to separate from other types of regular ur-
chins. Each interambulacral plate bears a
single, large, perforate primary tubercle
supporting a primary spine that is strongly
differentiated from the secondary spines
covering the rest of the test. Primaries are
many times larger than secondaries, and
much more robust. The shafts of adult ci-
daroid primary spines are unique not only
among all spines on a cidaroid, but also
among echinoids in general in completely
lacking an epithelial layer. These spines are
often invested with various epizoans such
as sponges, serpulids, bryozoans, and small
mollusks. Each primary is encircled by a
palisade of much shorter secondary spines
called scrobiculars that can be appressed to
the base of the primary spine.
226
In contrast to other regular echinoids, ci-
daroid ambulacra are very narrow relative
to the interambulacra. The ambulacral
plates are arranged in two simple columns,
each of which bears a single tube foot pore
(usually bipartite), and one to several small
secondary spines. The ambulacral series
continue to the mouth on the peristomial
membrane, and the number of peristomial
ambulacrals can be helpful in discriminat-
ing certain Antarctic taxa. The perignathic
girdle consists of interambulacral processes
called apophyses. The Aristotle’s lantern of
Antarctic species is typical for cidaroids,
and does not vary enough to form the
source of characters that could be used to
distinguish the taxa.
Although it is relatively easy to recog-
nize a cidaroid, a confounding feature of
the taxonomy within the group is conser-
vatism in plate architecture and overall
form of the test. However, the cidaroids
more than compensate for this conservatism
in diversity of primary spine morphology.
The taxa described in this paper exemplify
the fact that many cidaroid species cannot
be identified without information on the pri-
mary spines. Both scrobicular and non-
scrobicular secondary spines can also be
useful in diagnosing Antarctic cidaroids.
Cidaroids are epibenthic and inhabit var-
ious environments from sandy and stony
bottoms in littoral zones to gravels and
muds of sub-littoral regions. They are also
common in the deep-sea on muddy bottoms
of bathyal slopes or abyssal plains to 5000
meters, and they constitute a significant part
of the benthic community at these depths.
Many, if not all, of the Antarctic cidaroids
are direct developers, as suggested by the
fact that in almost every species, a subset
of adults presumed to be females have
greatly enlarged gonopores. Fell (1976) and
Lockhart et al. (1994) recorded that many
species also brood their young in and
around “‘marsupia”’ created by the more or
less sunken peristome, a feature of which
we make special note in the following de-
scriptions. Because the presence of brooded
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
young is correlated with enlarged gono-
pores, and because large gonopores are
known to signify females in other echi-
noids, we also use the presence of enlarged
gonopores and marsupia to infer gender in
adult specimens.
Upon occasion, primary spines of Ant-
arctic cidaroids also support a very poorly
understood parasite, Echinophyces mirabi-
lis Mortensen & Rosenvinge, 1909. The oc-
currence of this parasite is correlated with
some modifications of the infected echi-
noid’s morphology and a delay in the ap-
pearance and change in position of the gon-
opores. The mechanism by which a parasite
of a more or less biologically inert external
feature such as the cidaroid spine can affect
the expression of gonads and gonopores is
totally unknown and clearly deserves fur-
ther study. It is surprising that virtually no
research beyond the original description
and that of Mortensen & Rosenvinge
(1910) has been done on this unusual phe-
nomenon. Fell (1976), Jangoux (1987), and
Pearse & Cameron (1991) have summa-
rized what few data exist, finding that
Echinophyces is found only in Ctenocidaris
Mortensen, 1910 and Rhynchocidaris Mor-
tensen, 1909. We report on the results of
our as yet unsuccessful search for evidence
of this parasite in the new species, in the
hope that future studies might be able to use
even these negative data.
Order Cidaroida L. Agassiz, 1835
Family Cidaridae Gray, 1825
Genus Aporocidaris A. Agassiz & Clark,
1907
Diagnosis.—Ctenocidarine cidarids with a
large apical system which can be as much
as 75% of the horizontal diameter of the
test. Apical system often significantly
domed or convex. Mid-interambulacral re-
gions usually with a slightly depressed, na-
ked interradial suture.
Remarks.—Mortensen (1928) considered
four nominal species in the genus Aporo-
cidaris, but he emphasized the “‘exceeding-
VOLUME 113, NUMBER 1
ly slight” differences between 3 of them:
A. antarctica Mortensen, 1909, A. fragilis
A. Agassiz & Clark, 1907, and A. milleri
(A. Agassiz, 1898). Fell (1976) proposed
synonymy of these 3 species under A. mil-
leri but suggested a new species, A. eltan-
iana. In an unpublished manuscript, Chesh-
er recognized yet another species, A. usarpi.
We introduce the latter names to the pub-
lished literature for the first time.
Aporocidaris eltaniana, new species
Fig. 2
Aporocidaris eltaniana.—ex Fell, 1976:
211, figs. 7, 8j-k, name used in unpub-
lished thesis.
Diagnosis.—Apical system as much as
68% of test diameter. Aboral primary tu-
bercles large and markedly transversely
oval at ambitus. Aboral primary spines cy-
lindrical, 1 to 1.5 times test diameter, most-
ly smooth or with small bumps arranged in
longitudinal rows; tip blunt, or slightly con-
cave. Oral primary spines with conspicuous
thorns along lateral edges of shaft. Pre-
served specimens dark brown, primary
spines whitish, secondary spines beige to
light brown. Other characters as for Apo-
rocidaris.
Description.—The test is about half the
height of the horizontal diameter, but the
height of the apical system is quite variable
and can add as much as 20% to the overall
height of the coronal part of the test. The
largest recorded size of the specimens ex-
amined is about 45 mm in horizontal di-
ameter of the test. The holotype is 29.3 mm
in diameter, 16.9 mm high, has an apical
system diameter of 15.6 mm and a peri-
stome diameter of 12.5 mm.
The ambulacra are almost straight (Fig.
2A, B), particularly on the oral surface, and
their tuberculation is of the usual Aporoci-
daris pattern (Fig. 2E). There is no naked
area along the perradial suture. The inner
and outer pore of each podial pore pair are
divided by a well-developed ‘“‘bridge”’ of
stereom (Fig. 2E).
227
The interambulacra have about one more
plate in each series than A. milleri of similar
size. The crowding of the plates into the
relatively flat corona, and the size of the
tubercles between the ambitus and the peri-
stome cause the tubercles to be compressed
into horizontally oriented ovals (Fig. 2B).
The scrobicular tubercles are more than
twice the diameter of the other secondary
tubercles, and surround each primary tu-
bercle except where the oval outlines of the
primaries touch (Fig. 2D). The interradial
suture 1S conspicuously sunken in larger
specimens (Fig. 2D).
The apical system is large (about 60% of
the horizontal diameter on average), and al-
most flat or only slightly domed in most
specimens. However, a few exceptional in-
dividuals have a strongly arched apical re-
gion that can account for over a fifth of the
total height of the test. The genital plates
are large, with scattered secondary tuber-
cles. In males, the gonopores are small and
close to the outer edges of the genital plates
(Fig. 2A). In several of the specimens, the
gonopores are large, forming notches that
in extreme cases can extend part way along
the interradial suture separating the inter-
ambulacral plates adjacent to the genital.
Like Fell (1976), we have assumed these
specimens to be females. The gonopores
first appear in specimens as small as 16 mm
in test diameter. Ocular plates are only
about a third the area of the genitals, and
the ocular pores are surrounded by a cir-
cular ridge that seems to be most pro-
nounced in the males (Fig. 2A). Tubercu-
lation is not dense, and restricted to the cen-
tral parts of the plates over the majority of
the apex (Fig. 2A).
The peristome is just over 55% of the test
diameter in small specimens, but becomes
smaller in larger individuals (approximately
40% of the test diameter), relative to test
diameter. The peristome itself is somewhat
sunken around its periphery, notably in the
interambulacral regions. Seven to 8 pairs of
ambulacral plates continue onto the peri-
stome in larger specimens, and there are
228 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
==eee'
(NTR
AS
Oss
Wipe
f
fl
f iit A\\\ | @yY:
iter 5 ; Mgpttine bivee \
\\\
6) xe)
wee”
(A
a 3
Ss 2
S Zp
SSS
(y
Aboral plate pattern Oral plate pattern eh
Aboral }!1
primary (y
Scrobicular j
3 secondary
Side view of holotype spine
Non-scrobicular }
secondary §
Fig. 2. Aporocidaris eltaniana new species. C drawn from holotype (NMNH E48122); A, B, D—I drawn
from paratypes in lot NMNH E11161. For A and B: mouth, anus, and gonopores in solid black; podial pores
omitted; secondary tubercles shown only for single genital, ocular, and periproctal plates, but omitted elsewhere,
including coronal plates.
VOLUME 113, NUMBER |
only 2 or 3 small scales in each interradial
region (Fig. 2B).
The longest aboral primary spines found
(on the almost intact specimens in NMNH
E11161) are approximately 1.5 times the
test diameter and rod-like with blunt or
slightly concave tips and large milled rings
(Fig. 2F). The surface of the shaft is smooth
proximally, but bears small, longitudinally
aligned, blunt spinules for more than 80%
of its length out to the tip (Fig. 2F). The
flat areas between the longitudinal rows are
covered with a variously developed fibrous
and anastomosing calcareous meshwork.
The oral primaries are strongly curved to-
wards the peristome, and bear well devel-
oped thorns that project laterally, but be-
come smaller towards the tip of the spine.
There is often another row of shorter thorns
along the distal part of the shaft (Fig. 2G).
The more or less cylindrical scrobicular
spines are blunt, but not club-shaped (Fig.
2H). The non-scrobicular secondaries are
much shorter than the scrobicular spines
and sometimes slightly curved near the al-
most imperceptibly swollen tip (Fig. 21).
The small globiferous pedicellariae are
rare, but appear simply to be half-sized ver-
sions of the larger type. The stem of a glob-
iferous pedicellaria is thick, and straight.
Neither the valves nor stem are otherwise
distinctive among Aporocidaris species
(Mortensen 1928).
Types.—Holotype NMNH E48122, ex-
tracted from NMNH E11161, R/V Eltanin
Cruise 6, Station 432 (Fig. 2C). Paratypes
NMNH E11161 (3 dry specimens, one dis-
sected lantern), E11188 (17 dry specimens,
including 3 less than 10 mm in horizontal
test diameter), E11212 (dry spines only),
E11290 (more than 60 dry specimens),
E14597 (2 specimens greater than 28 mm
in test diameter, the largest of which is a
female with more than 12 young urchins
brooded on the peristome, 1 specimen 14.3
mm in horizontal test diameter, and 16
specimens less than 10 mm in test diameter,
all in alcohol), MCZ 8406 (1 dry speci-
men). The holotype was chosen because it
229
retained significant spine cover (Fig. 2C).
The paratypes in NMNH E11161 were used
to make the drawings of plate architecture
so as not to disturb what spination remains
on the holotype.
Etymology.—Named for U.S. National
Science Foundation R/V Eltanin. The spe-
cies name was first coined in an unpub-
lished doctoral thesis by Fell (1976), and
we introduce the name here in recognition
of that first usage.
Distribution.—A. eltaniana is known
only from the type series, which was col-
lected from two R/V Eltanin stations
(Cruise 4, Station 138; Cruise 6, Station
432) at two distinct localities, both in the
region of Livingston Island in the South
Shetland Islands, between 884 and 1437 m
depth (Fig. 1).
Biology.—The gonopores in females
may reach 1.8 mm in diameter. One female
(NMNH E14597), 43.4 mm in horizontal
test diameter, had more than a dozen juve-
nile urchins in the marsupium along the
sunken edge of the peristomial margin. An
exact count of the juveniles was impossible
without damaging the adult specimen. Most
of the young were obscured by oral primary
and secondary spines that folded over to
screen the deepest parts of the marsupium
in the interambulacral areas proximal to the
edge of the coronal plates. One of the larg-
est of the juveniles was 2.2 mm in horizon-
tal test diameter, and all of the brooded
young possessed primary spines, the lon-
gest of which were approximately 2.0 mm
long. These observations suggest that ju-
veniles of A. eltaniana develop directly
from relatively large eggs which are moved
into, and then develop in a peristomial mar-
supium. None of the specimens appear to
be infected by Echinophyces. Nothing is
known of the preferred bottom type, but the
gut contents of one specimen are made up
of light gray silt containing small fragments
of thin mollusk shells.
Remarks.—The greatly enlarged apical
system and the naked, sunken area along
the interradius indicate that the new species
230
is an Aporocidaris. A. eltaniana is most
similar to A. milleri, but is distinct from all
Aporocidaris in possessing strongly devel-
oped thorns on the oral primary spines. The
dramatic difference between the relatively
smooth, cylindrical aboral primaries and the
thorny oral primaries is unmatched in the
genus, if not within the Antarctic cidaroids
in general.
Aporocidaris usarpi, new species
Fig. 3
Diagnosis.—Apical system as much as
70% of test diameter in adults. Aboral pri-
mary spines cylindrical, 1 to 1.5 times test
diameter, with small, irregularly distributed
or only slightly aligned bumps distally.
Neck of aboral primaries glossy, porcella-
neous, and markedly swollen in larger spec-
imens. Oral primary spines flattened with
lateral, diametrically opposed, serrated
keels that are widest just distal to neck and
taper towards spine tip. Scrobicular spines
fine, small, closely spaced, and not strongly
differentiated from the non-scrobicular
spines. Both types of secondary spines
slightly club-shaped. Preserved specimens
beige, primary spines white, secondary
spines light beige. Other characters as for
Aporocidaris.
Description.—The test is only moderate-
ly flattened. The height of small specimens
is Just over 50% of the horizontal diameter,
but in larger specimens, the height is greater
(up to 75% of the horizontal diameter), al-
most 15% of which can be in the form of
the domed apical system (Fig. 3C). The
largest specimen is just over 40 mm in hor-
izontal diameter. The holotype is 35.2 mm
in diameter, 23.4 mm high, has an apical
system diameter of 23.0 mm and a peri-
stome diameter of 15.2 mm.
The ambulacra are almost straight (Fig.
3A, B), particularly on the oral surface, and
their tuberculation is of the usual Aporoci-
daris pattern, with smaller secondary tuber-
cles adjacent to the perradial suture, and
Slightly larger ones just distal to those (Fig.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
3E). There is no naked region along the per-
radius. In the holotype and the largest of the
paratypes (NMNH E11059), the “bridge”
between the pore pairs on each ambulacral
plate is incomplete so that the pores are con-
nected via a constricted passage between the
pores (Fig. 3E). This condition is rare, if not
unique among Antarctic cidaroids.
The interambulacral plates are high. Ab-
orally, the primary tubercles do not occupy
the entire height of the plate so that the ab-
oral primary tubercles retain their round
outlines. However, adjacent to the peri-
stome, the tubercles are more crowded, and
tend to compress slightly into transverse
ovals (Fig. 3B). The scrobicular tubercles
are not strongly differentiated in size from
the non-scrobicular secondaries, which are
small and closely spaced everywhere out-
side the primary tubercle except along the
interradius, where tubercles are lacking
(Fig. 3D). This distinct naked area is adja-
cent to each interradial suture and occupied
by a shallow furrow that, in larger speci-
mens, sends branches circumferentially
along sutures separating plates in each of
the two interambulacral columns (Fig. 3D).
The apical system is large, ranging from
just over 50% of the test diameter in juve-
niles to over 70% of the diameter in the
largest adults. In the juveniles, the apical
region tends to be almost flat. It is distinctly
domed in adults, with considerable irregular
inflation of the centers of some of the larger
apical plates (particularly the genitals), es-
pecially in the specimen identified as a fe-
male. The genital plates are large relative
to the other plates in the apical system. In
the largest paratype, which appears to be a
female, the gonopores can be as much as
2.4 mm in diameter and in some cases oc-
cupy shallow clefts along the interradial su-
ture (Fig. 3A). The gonopores are present
in the holotype, which appears to be a male,
but are not evident in the larger of the 2
small paratypes in NMNH E11059, which
is 13.9 mm in diameter. Ocular plates are
only about a quarter the area of the genitals,
and the ocular pores are surrounded by a
VOLUME 113, NUMBER 1
prominent circular ridge (Fig. 3A). The tu-
berculation on both oculars and genitals is
fine and dense, and tends to cover the entire
plate surface almost to the suture (Fig. 3A).
The peristome is sunken around its pe-
riphery, and in the region of half the test
diameter. Each perradial region bears 8 or
9 plates in each ambulacral column, and
there are two relatively large scales in each
interradial portion of the membrane (Fig.
3D).
The longest aboral primary spines are ap-
proximately 1 to 1.5 times the horizontal
diameter of the test, although even the best
examples are worn or broken so that the
nature of the tips cannot be ascertained.
Distal to the neck, the shaft is cylindrical
and rod-like, and invested with irregularly
spaced to only partially aligned, short, blunt
spinules or bumps that arise from _ the
smooth, porcellaneous surface. The neck is
most peculiar in that it is greatly swollen in
primary spines of larger specimens, and
completely unadorned with spinules. The
surface of the swelling is smooth, porcel-
laneous and glossy, and increases the di-
ameter of the spine by a factor of 2 at its
widest point (Fig. 3F). In juvenile speci-
mens, the neck is also porcellaneous but
only slightly swollen, suggesting that the
vase-like shape of the swelling becomes
more prominent with age. The oral primary
spines are only slightly curved towards the
peristome, and bear diametrically opposing
flanges or keels along the lateral sides of
the shaft. The edges of these keels are finely
serrated, and the keels themselves are wid-
est proximally, tapering towards the rela-
tively blunt spine tip to give the entire spine
a dagger-like appearance (Fig. 3G). The
surface of the oral primaries is smooth and
shiny, but not as glossy as the aboral pri-
maries. The almost cylindrical scrobicular
spines are blunt and very slightly swollen
towards the tip (Fig. 3H). Scrobicular sec-
ondaries are only slightly longer than the
non-scrobicular secondary spines (Fig. 3I).
Both types of secondaries are club-shaped.
The valves of the large globiferous ped-
2351
icellariae are less than 1 mm in length, and
very similar to those of other Aporocidaris
(Mortensen 1928). The smaller globiferous
were not detected.
Types.—Holotype NMNH E11134, R/V
Eltanin Cruise 14, Station 1212 (Fig. 3C,
F-I). Paratypes NMNH E11059 (3 dry
specimens, one dissected lantern, and one
almost perfect, loose spine), E14603 (1
specimen in alcohol). The holotype was
chosen because it retained some spines
(Fig. 3C). Plate architecture was drawn
from one of the largest of the paratypes in
NMNH E11059 to avoid further damage to
the holotype.
Etymology.—The name “‘A. usarpi’’ was
originally used by Dr. Richard Chesher in
his identifications of USARP material now
housed the NMNH (see above), and it is
clear from notes placed with that material
that he intended to publish the species in a
manuscript that never saw press. We retain
Chesher’s suggested name in honor of his
recognition of the distinctiveness of this
taxon.
Distribution.—A. usarpi is known only
from a single R/V Eltanin locality (Cruise
14, station 1212) from a mid-ocean point
far to the southeast of New Zealand and
north of the Ross Sea (Fig. 1) at a depth of
between 3678 and 3935 m.
Biology.—Nothing is known of the hab-
itat preferences of A. usarpi, and appropri-
ate specimens could not be sacrificed for
gut content observations. There is some
sexual dimorphism. The putative female
has greatly enlarged gonopores (up to 2.4
mm in diameter) relative to the putative
male, suggesting direct development in this
species. None of the specimens had young
retained on the test, although the peristo-
mial edge is slightly sunken in such a way
as to suggest the same type of marsupia
seen in other species bearing young on the
peristome. None of the specimens show ob-
vious signs of Echinophyces infection, but
the paucity and condition of the material
prevent us from ruling out the occurrence
of the parasite in A. usarpi.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
[SN
Aboral plate fr C7 ‘
pattern by i ‘ ES 6 ©
0° et e 7 NN \
EZ @y
ay
rd LE
: [Pr bo%
((uauatte
1D)
ies
ONS
ree
S
o \ fe
Side view of G a H primnanyt
holotype Oral ("8 spine}:
< primary (. 23 Scrobicular
pe spine(® #4 secondary
‘} a spine
Non-
scrobicular
secondary
spine
Ambulacrum at ambitus
Fig. 3. Aporocidaris usarpi new species. C, F—I drawn from holotype (NMNH E11134); A, B, D, E drawn
from paratypes in lot NMNH E11059. Conventions for A and B as in Fig. 2.
VOLUME 113, NUMBER 1
Remarks.—The greatly enlarged apical
system and the naked, sunken area along
the interradius indicate that the new species
is an Aporocidaris. The species is readily
distinguished from all other Aporocidaris
by the peculiar swelling of the neck in the
primary spines, and the extreme glossiness
of both this swelling, and the shaft of the
spines between the distal spinules. The ab-
sence of stereom bridges separating the
members of the podial pore pairs has not
been previously reported for any ctenoci-
darine taxa, let alone other Aporocidaris.
Both the holotype and the largest of the
paratypes exhibit this feature, but the two
smallest specimens (below 14 mm horizon-
tal diameter) have clearly divided pores,
suggesting that the unification of the pores
progresses with ontogeny, and is a derived
feature of adult A. usarpi.
Genus Notocidaris Mortensen, 1909
Diagnosis.—Oral primary spines spear-
or dagger-shaped. Aboral primaries rod-like
and cylindrical to flattened and spatulate,
with large lateral expansions distally.
Spines smooth or spiny, but when present,
spinules restricted to proximal part of shaft.
Proximal surface of spines occasionally
with coating of anastomosing hairs. Scrob-
icular secondary spines simple and slender
(but not as fine or densely distributed as in
Aporocidaris), not distinct in shape from
other secondary spines. Interradial sutures
naked, but not sunken. Apical system about
60% of horizontal diameter.
Remarks.—Mortensen (1928) listed 4
species in the genus: N. gaussensis Morten-
sen, 1909, N. hastata Mortensen, 1909, N.
mortenseni (Koehler, 1900), and N. platya-
cantha (H. L. Clark, 1925). Another spe-
cies, N. remigera Mortensen, 1950 was de-
scribed after Mortensen’s monograph of the
cidaroids was published. We do not consid-
er the entity N. platyacantha var. contracta
Koehler, 1926 to be a valid taxon. There-
fore, we attribute 5 full species to Notoci-
daris, all from Antarctic and Subantarctic
233
waters, and all distinguished by features of
the aboral primary spines. It should be not-
ed that the range of variation recorded for
each species can be extremely large and
spine characteristics can overlap for species
such as N. remigera and N. mortenseni.
Therefore, biogeographic and bathymetric
data, as well as other parts of the descrip-
tions must also be used to make positive
determinations.
Notocidaris lanceolata, new species
Fig. 4
Notocidaris lanceolata.—ex Fell, 1976:
195; figs. 7, 8f-g, name used in unpub-
lished thesis
Diagnosis.—Aboral primary spines
coarsely thorned at their base, with thorns
arranged irregularly, and becoming greatly
reduced in size and aligned towards tip.
Spines keeled in two orthogonal planes in
cross section, yielding lance-like appear-
ance, spine tapering slightly towards rela-
tively blunt tip. Longest spines about 2
times horizontal diameter of test in most
specimens. Apical system varying from flat
to arched. Test of preserved specimens
beige to ochre, primary spines beige with
shiny purplish-beige neck, secondary spines
beige. Other characters as for Notocidaris.
Description.—The largest recorded hori-
zontal test diameter is 36 mm. The height
is between 50% and 60% of the test diam-
eter. The holotype is 32.5 mm in horizontal
diameter and 19.4 mm high.
The ambulacra are straight on the oral
surface, with some slight sinuousness ab-
orally (Fig. 4A, B). There are one or two
secondary tubercles perradial to the podial
pores, and two or three much smaller sec-
ondary tubercles next to the perradial suture
(Fig. 4E). There is no naked region along
the perradius. The inner and outer pore of
each podial pore pair are divided by a well-
developed “‘bridge’’ of stereom (Fig. 4E).
The aboral interambulacral plates are
high and the primary tubercles are circular.
Orally, the plates are not as high and the
234
tubercles are compressed into transverse
ovals (Fig. 4B). There are 7 or 8 plates in
each column in a specimen about 50 mm in
diameter. The scrobicular tubercles are in
general only slightly differentiated from the
non-scrobicular secondaries, which are
abundant and closely spaced except along
the interradial suture, where tubercles are
lacking (Fig. 4D). There is no sunken area
along the interradial suture.
The apical system is usually about 60%
of the test diameter in larger specimens, and
can range from being flat, as in the holo-
type, to domed. The gonopores are restrict-
ed to the genital plates and do not signifi-
cantly invade the adjoining interambulacra.
The females have enlarged gonopores. The
oculars are large and circumferentially
elongated so that they are much wider than
they are high, and about half the surface
area of the genital plates (Fig. 4A). The oc-
ular pore is encircled by a prominent ridge
and the tuberculation is fine and evenly dis-
tributed across each plate in the apical sys-
tem except directly adjacent to the sutures
(Fig. 4A).
The peristome is only slightly sunken at
its periphery, and approximately 45% of the
test diameter. There are 7 or 8 plates in each
ambulacral column on the peristome of
larger specimens, and as many as 5 small,
irregularly shaped scales in each interradial
portion of the membrane (Fig. 4B).
The ambital and aboral primary spines
can be up to 2 times the diameter of the test
and tapered. Several complete examples re-
main attached to the holotype (Fig. 4C).
Near the base, the spine is adorned with
irregularly arranged, prominent thorns and
serrations which diminish sharply in size
about 30% of the spine’s length away from
the base, giving way to strongly aligned,
distal rows of spinules (Fig. 4F). Although
the cross-section of the spine is broadly cir-
cular, usually there are also well-developed
keels set almost orthogonally to each other
along the distal part of the spine. The de-
gree to which these keels are developed
varies among specimens and even on an in-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
dividual, but the basal thorns seem to be a
constant feature. Basally, the spine can bear
a surface coat of anastomosing hairs similar
to that seen in some Aporocidaris. The oral
primaries are differ from the aboral prima-
ries in lacking strongly developed keels ex-
cept laterally, where they can also be
strongly serrated (Fig. 4G), particularly in
specimens with thorny aboral primaries.
The oral primaries adjacent to the peristome
are very small and dagger- or arrowhead-
shaped, with somewhat less distinctly ser-
rated keels. In specimens with reduced
keels on the aboral primaries, the oral pri-
maries are less dagger-shaped, and more
oval in outline. The scrobicular spines are
slightly pointed (Fig. 4H). The non-scrobi-
cular secondary spines are smaller, partic-
ularly in the ambulacra (Fig. 41).
As in most ctenocidarines, the globifer-
ous pedicellariae come in two sizes, both of
which are quite long and slender in N. Jan-
ceolata, though not diagnostically so. The
valves can be just over 1 mm long in the
larger form.
Types.—Holotype MCZ 8336, R/V EI-
tanin Cruise 32, Station 2110 (Fig. 4C).
Paratypes NMNH E21865 (3 dry speci-
mens), E21866 (1 dry specimen), E22004
(1 specimen in alcohol), E22005 (5 speci-
mens in alcohol), E22006 (3 specimens in
alcohol). We decided to retain the holotype
designated in Fell (1976) to avoid confu-
sion, and also because the specimen has ex-
emplary spine cover (Fig. 4C). Drawings of
spines and plate architecture were made
from the paratypes to avoid damaging the
holotype.
Etymology.—In his thesis, Fell (1976) at-
tributed the name to Richard Chesher, who
recognized the distinctiveness of the new
taxon in a manuscript that was never pub-
lished. Chesher based the name on the
keeled, lanceolate spines.
Distribution.—Known from 7 R/V El-
tanin stations (Cruise 27, Stations 1867,
1926, 1929; Cruise 32, Stations 2002, 2108,
2110, 2129) at three distinct localities in the
VOLUME 113, NUMBER 1 to
A a F
Aboral plate LEZ = Aboral /',
pattern primary {'f
C
Side view
of holotype fj
©
] nA
es
\Sy yet
s {| |
NG GAS
~ GY NV, Ss H fi
We... ee Scrobicular | |
secondary 1
spine
2! Interambulacrum at ambitus ea ern at ainbhGs
Fig. 4. Notocidaris lanceolata new species. C drawn from holotype (MCZ 8336); A, B, D, E drawn from
paratypes in lot NMNH E21865; F—I drawn from paratype (NMHH E 21866). Conventions for A and B as in
Big: 2.
236
Ross Sea between 2005 and 2421 m (Fig.
1):
Biology.—The spines seem remarkably
free of epizoans. The preferred habitat is
unknown. Some specimens, which could be
females, have enlarged gonopores, but no
broods have been observed. No evidence of
Echinophyces infection could be detected.
Remarks.—As noted by Fell (1976:197),
the species was based on a specimen res-
cued by Dr. Merrill Foster from ‘material
intended for disposal overboard”’ and later
turned over to Chesher, who selected it as
a potential type. The holotype, in having
primaries that are thorny and strongly lan-
ceolate, is close to one end of a range that
includes forms in which the keels can be
almost absent. As Fell (1976:197) indicat-
ed, “‘were it not for the range of interme-
diates between fluteless [unkeeled] and ful-
ly fluted [keeled] specimens obtained at two
stations, one would not believe they were
of a single species”’. Typical N. lanceolata
may be most easily confused with particu-
larly thorny N. hastata, especially since
they inhabit the same environments, while
variants of N. lanceolata with greatly re-
duced keels on the spines may be confused
with N. gaussensis. In addition, the surface
coating of anastomosing hairs between the
thorns and keels near the base of the spine
can cause some confusion with Ctenocidar-
is, particularly if this feature alone is used
to sort material. Because N. lanceolata oc-
curs, on average, almost 1000 m deeper
than N. gaussensis and virtually all Cteno-
cidaris, available depth data should help to
provide initial clues in identification.
Acknowledgments
We would like to thank Cynthia Ahearn
(National Museum of Natural History) for
her patience and helpfulness throughout our
endless inquiries and loan requests and
Chris Mah of the California Academy or
Sciences (California Academy of Sciences)
for digging out distant data. Fred Collier
(Museum of Comparative Zoology) provid-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ed invaluable information concerning the
type of N. lanceolata. We are also grateful
for the editorial comments of John Pearse
and Dave Pawson. Mooi’s research was
supported by an American Philosophical
Society General Research grant. This paper
is a contribution of the theme ‘‘Signal mor-
phologique de Evolution” of the UMR
CNRS 5561 “‘Biogéosciences.”’
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Smith, A. B. 1984. Echinoid palaeobiology. Allen and
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, & C. W. Wright. 1988. British Cretaceous
echinoids. Part 1, General introduction and Ci-
daroida. Monograph of the Palzontographical
Society, London—141:1-—101.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):238—248. 2000.
A new species of Pristigaster, with comments on the
genus and redescription of P. cayana
(Teleostei: Clupeomorpha: Pristigasteridae)
Naércio A. Menezes and Mario C. C. de Pinna
(NAM) Museu de Zoologia, Universidade de Sao Paulo, Av. Nazaré 481,
Sao Paulo-SP 04263-000, Brazil; (MCCdeP) Departamento de Zoologia,
Universidade de Sao Paulo, Caixa Postal 11461, Sao Paulo-SP 05422-970, Brazil
Abstract.—A new species of the hitherto monotypic genus Pristigaster (Clu-
peiformes: Pristigasteridae) is described for the Amazon basin. Pristigaster
whiteheadi, new species, is distinguished from its only congener, P. cayana,
by the presence of pelvic fins; the lack of caudal-fin filaments; the different
angle between the predorsal bones and the vertebral column; the presence of
36—39 vertical scale rows (40—47 in P. cayana); and the presence of 18—20
horizontal scale rows (21—26 in P. cayana). Pristigaster cayana is redescribed,
and its occurrence in French Guyana is questioned.
Resumo.—Uma nova espécie do género até entao monotipico Pristigaster
(Clupeiformes: Pristigasteridae) é descrita para a bacia Amaz6nica. Pristigaster
whiteheadi, espécie nova, distingue-se de P. cayana pela presenga de nadad-
eiras pélvicas; auséncia de filamentos nos lobos da nadadeira caudal; o 4ngulo
diferente entre os ossos pré-dorsais e a coluna vertebral; a presenga de 36—39
fileiras verticais de escamas (40—47 em P. cayana); e a presenga de 18—20
fileiras horizontais de escamas (21—26 em P. cayana). Pristigaster cayana é
redescrita e sua ocorréncia na Guiana Francesa é questionada.
The Clupeomorpha is a highly diverse
group of teleosts, containing over 350 re-
cent and over 150 fossil species (Grande
1985, Nelson 1994). The group is one of
the most important fisheries resources
worldwide, and their phylogenetic relation-
ships within teleosts have been a matter of
intense debate in recent years. For the past
two decades, following the original sugges-
tion by Patterson & Rosen (1977), Clupeo-
morphs were placed as the sister group to
the Euteleostei. More recently, Clupemor-
pha have been proposed as sister group to
Ostariophysi, a hypothesis supported by
molecular data (Van Le et al. 1993, Patter-
son 1994) and morphological characters
(Lecointre & Nelson 1996, Johnson & Pat-
terson 1996, Arratia 1997).
Clupeomorphs are a demonstrably mono-
phyletic group (Grande 1985) including
basal fossils as the extinct genera Diplo-
mystus and Armigatus, and the extinct order
Ellimmichthyiformes. Recent clupeo-
morphs are all in the order Clupeiformes,
itself divided into suborders Denticipitoidei
(with a single species from African fresh-
waters) and Clupeoidei (all other recent clu-
peiforms). Clupeoids comprise three super-
families: Engrauloidea (with a single fami-
ly, Engraulididae), Clupeoidea (with fami-
lies Chirocentridae and Clupeidae) and
Pristigasteroidea (with families Pristigaster-
idae and Pellonidae).
The pristigasteroid family Pellonidae in-
cludes the central and South American gen-
era Chirocentrodon, Neoopisthopterus, Pel-
lona, and Pliosteostoma. The Pristigasteri-
dae, in turn, comprises the Central and
VOLUME 113, NUMBER 1
South American Odontognathus and Pris-
tigaster, the South American and Indo-Pa-
cific Opisthopterus and the Indo-Pacific Ra-
conda. The genus /lisha could not be dem-
onstrated monophyletic by Grande (1985),
most of its species forming a polytomy at
the base of Pristigasteroidea.
Pristigaster is the most peculiarly-shaped
of all pristigasteroids, with an extremely
deep body resembling characiforms of the
genera Gasteropelecus and Thoracocharax.
So far a single species, P. cayana, is rec-
ognized in the genus. All other proposed
names have been shown to be either invalid
or junior synonyms of that species (White-
head 1973, 1985). However, Whitehead
(1985) suggested that a second species
might exist. Stimulated by Whitehead’s
original suggestion, we undertook a de-
tailed examination of available material of
Pristigaster, and concluded that indeed
there is a second diagnosable species in the
genus, still undescribed. In this paper, we
formally name and diagnose the new spe-
cies and redescribe P. cayana.
Methods and materials.
Morphometric measurements were all
point-to-point, taken with calipers, recorded
to the nearest 0.1 mm and expressed as per-
centages of standard length, except for sub-
units of the head, expressed as percentages
of head length. Counts and measurements
were made on the left side of the speci-
mens, whenever possible, according to
Whitehead (1985), except for horizontal
rows of scales (counted between dorsal-fin
origin and anal-fin origin), vertical rows of
scales (counted from origin of pectoral fin
to caudal base), and scales around caudal
peduncle (number of horizontal scale rows).
Principal caudal-fin rays included all
branched rays plus one unbranched ray in
each lobe. Counts for each lobe, upper first,
are separated by a slash. Vertebral counts
were taken from radiographs and cleared
and stained specimens and the terminal
“half centrum” is included. Specimens
239
were dissected to determine sex by an in-
cision on the right side of the abdomen to
expose the gonads. Tooth counts include
sockets in cases where the actual tooth has
fallen off. Within the meristic information
given here, figures for holotype are provid-
ed in parentheses.
Figures associated with specimen lists in
species descriptions are, first, number of
specimens examined in respective lot, and
second, range of SLs in mm.
Specimens cleared and counterstained for
bone and cartilage were prepared by a mod-
ified version of the method of Taylor & Van
Dyke (1985). Descriptive accounts follow
the general organization in Whitehead &
Teugels (1985), the most complete anatom-
ical survey of a clupeomorph available to
date. Synonymic lists include only those
references in which the species referred to
can be reliably identified as either P. cay-
ana or P. whiteheadi.
Specimens examined in this work are
deposited in the following institutions:
AMNH, American Museum of Natural
History, New York; BMNH, The Natural
History Museum, London; FMNH, Field
Museum of Natural History, Chicago;
INPA, Instituto Nacional de Pesquisas da
Amazonia, Manaus; MZUSP, Museu de
Zoologia, Universidade de Sao Paulo, Sao
Paulo.
Pristigaster whiteheadi, new species
Fisssf) 2B
Pristigaster cayana (not Cuvier); White-
head, 1985:301 (in part, only specimens
with pelvic fins); Whitehead & Bauchot,
1985:24 (in part, only specimens with
pelvic fins); Stewart, Barriga & Ibarra,
1987:21 (specimen examined).
Holotype.-—MZUSP 52963 (female, 83.4
mm SL). BRAZIL: Amapa, Rio Araguari,
Ferreira Gomes, collected by M. Goulding,
January—February, 1984.
Paratypes.—Brazil: MZUSP 30341 (2,
76.5—83.4), same data as holotype. Ama-
zonas; MZUSP 11391 (8, 57.5—67.7, 2 of
240
which cleared and counterstained), AMNH
227329: (15'52:2); USNIM351306°G1; 6220);
FMNH 107783 (1, 56.5), Rio I¢a, Santo
Anténio do I¢a4; MZUSP 11392-393 (2,
69.4—73.0), Rio Solim6es, above mouth of
Jutai; MZUSP 11394—403 (10, 43.5—76.2),
Rio Solimdes, Fonte Boa; MZUSP 27597
(1, 62.00, Rio Solimdes, Municipio de Ben-
jamin Constant; MZUSP 18694 (3, 29.4—
40.0), Rio Solimdes, Lago Janauaca and vi-
cinity; MZUSP 52950 (2, 22.0—25.0), Rio
SolimGes, 3°10'57”S, 67°56'31"W; MZUSP
6600 (1, 70.0), Lago Manacapuru; MZUSP
18512 (1, 67.0), mouth of Rio Ituxi;
MZUSP 18516 (1, 70.0), mouth of the Pa-
cia; INPA 8555 (21, 18.0—66.6, 3 of which
cleared and stained), Parana do Tapura, near
mouth of Rio Madeira; MZUSP 6220 (1,
84.0), Rio Negro, Igarapé Jaraqui, above
Manaus; MZUSP 52951 (1, 39.0), Rio
Jauaperi, 1°34'54"S, 61°28'48"W; MZUSP
52952 "(25 '57-7 ‘and 167.0) R10" Neero;
1°33'48”"S, 61°33'02"W; MZUSP 49597 (3,
28.3—37.3), Rio Acre, above Boca do Acre;
MZUSP 7625 (1, 67.0), Rio Amazonas, Pa-
rana do Mocambo, above Parintins; BMNH
1897.12.1.197—-199 (3, 62.8—-65.0), Rio Ju-
rua; MZUSP 52949 (21, 47.0—86.0), Rio
Japurad, Parana do Japurd, 3°09'12"S,
64°46'54"W; MZUSP 52962 (1, 63.3), Rio
Amazonas, 1°54’S, 55°31’W; MZUSP
52948 (5, 35.5—76.0), Rio Madeira, below
Nova Olinda; MZUSP 52958 (1, 36.0), Rio
Madeira, 3°33'37"S, 58°59'49"W; MZUSP
52957. (1, 34.0)23° 3348S. 585957 Ww
MZWSP »529592 Gs 4120)53 29) 2S,
538°5.1°338" W, .MZUSP.. 52960. (1... 30:5):
3°26'44"S, 58°49’49"W, MZUSP 52961 (4,
30.5—41.0), 3°33’S, 58°55'W, Rio Madeira,
Parana do Urucurituba. Roraima: MZUSP
11404 (1, 33.0), Rio Branco, 20 kilometers
below Boiacu. Paraé: MZUSP 52953 (1,
28.0); 1535" S,.5271 1 WAMZUSP#52954. (7,
50.0—55.0), 1°27’S, 52°03’W, Rio Amazon-
as, Furo do Urucuricaia; MZUSP 5493 (1,
85.0), Rio Trombetas, Oriximina; MZUSP
529558(1;. 53.7),4 1529'S 52°35 Wa MZUSP
52956 (8, 22.3—66.5), 1°36’S, 52°12'W, Rio
Amazonas, below Rio Xingu. ECUADOR.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
FMNH 101946 (1, 89.0) 0°49’S, 75°31'W,
Rio Tiputini, near mouth in Rio Napo and
Quebradas).
Diagnosis.—Distinguished from its only
congener, P. cayana, by the following fea-
tures: 1—presence of pelvic fins; 2—ab-
sence of filaments on upper and lower
lobes of caudal fin; 3—-vertical scale rows
36-39 (40—47 in P. cayana); 4—horizon-
tal scale rows 18—20 (21-26 in P. cayana);
5—supraneurals (predorsal bones) gradu-
ally less sloped posteriorly, posterior one
nearly perpendicular to vertebral column
(supraneurals all equally sloped in P. cay-
ana). Most specimens of the new species
can also be distinguished from P. cayana
by lower gill raker counts (18—21, versus
21-25 in P. cayana) and by lower anal-fin
ray counts (41—48 versus 44—53 in P. cay-
ana).
Description.—Meristic and morphomet-
ric data are presented in Tables 1 and 2. For
a general aspect of the fish, refer to Fig. 1.
Body highly compressed, ventral profile of
body extremely expanded and convex, its
anterior region (at isthmus) almost perpen-
dicular to longitudinal axis of fish. Five
protruding predorsal supraneurals strongly
inclined anteriorly. The entire abdominal
region, from isthmus to anal-fin origin, bor-
dered by a series of 29-34 (holotype 31)
abdominal scutes, gradually more promi-
nent posteriorly. Scutes anterior to vertical
through pectoral-fin base mostly imbedded
in soft tissue, those posterior to that point
protruding markedly beyond abdominal
profile resembling a series of translucent
hooks. Pelvic-fin origin usually over 25th
scute (as in holotype), rarely over 24th or
26th scutes.
Snout blunt, always shorter than orbital
diameter. Mouth subterminal and turned
dorsally, its lower jaw protruding beyond
upper. Maxilla extending slightly posterior
to vertical through anterior margin of eye.
Teeth conical, minute, disposed in a single
irregular row in both jaws and highly var-
iable in number, becoming more numerous
with growth. Premaxillary teeth 8 (in spec-
VOLUME 113, NUMBER 1
Table 1.—Selected meristic features of Pristigaster cayana and P. whiteheadi.
Pristigaster cayana Pristigaster whiteheadi
n Range Mean SD n Holotype Range Mean SD
Branched dorsal-fin rays 79 12-13 12.177 0.384 95 12 11-13 12.053 0.305
Branched pectoral-fin rays 79 10-12 10.848 0.622 95 10 oI) 710.3537 0,522
Branched pelvic-fin rays 95 3 3-4 3.137 0.346
Branched anal-fin rays 73 44-53 48.096 2.076 93 44 41-48 44.258 1.436
Gill rakers TT Y23=25+° 22,208 - 0:8468 95 Ps 18-21 19.895 0.722
Horizontal rows of scales 55 21-26 23.182 1.090 a, — 18-20 19.114 0.631
Vertical rows of scales 39 40-47 43.077 1.797 18 — 36-39 38.222 0.878
Scales around caudal peduncle 18 15-16 ~13,833- 0.383 9 — 13-15", 14.556. 0.726
Premaxillary teeth 58 9-23 16.862 3.322 78 14 8-18 11.962 1.970
Maxillary teeth 62 18-80 53.871 14.140 84 63 14-64 40.000 11.600
Dentary teeth 49 5-14 9.408 2.188 53 5-9 ye lw: 1.301
Ventral scutes Ta. 30535 32,013 IEP | 92 31 29-34 31.065 0.849
Vertebrae 7 42-44 43.428 0.728 | — 43-44 43.286 0.488
imens <44 mm SL) to 18 in larger speci-
mens (holotype 14). Maxillary teeth 14 (in
22 mm SL specimen) to 64 (in 79 mm SL
specimen) (holotype 63). Dentary teeth 6
(at 43 mm SL) to 9 in larger specimens
(holotype 9). Eyes very large, round in ex-
ternal aspect. Pupil extremely large rela-
tive to eye size, its diameter almost 60%
that of orbit. Adipose eyelid weakly de-
veloped in young, but covering most of iris
in adults.
Gill cover rounded in profile, with con-
tinuous membranous margin concealing a
marked depression on posterior margin of
opercle. Dorsal limit of gill opening slightly
ventral to horizontal at dorsal margin of
eye. Gill rakers slender, closely set and con-
spicuous, their number increasing with
growth, ranging from 18 (at 22 mm SL) to
21 (at 83 mm SL) (holotype 21) on lower
part of first branchial arch and from 8 to 12
(holotype 11) on upper part.
Table 2.—Morphometrics of Pristigaster cayana and P. whiteheadi. Standard length is expressed in mm;
measurements | to 12 are proportions of standard length; 13 to 16 are proportions of head length.
Pristigaster cayana
Character n Range Mean
Standard length 79 =—-.23.5-142.0 66.68
1. Body depth 78 0.38-0.65 0.550
2. Predorsal length 79 ~=0..39-0.48 0.421
3. Prepectoral length [Bie o0.23-031 0.271
4. Prepelvic length
5. Preanal length 78 0.57-0.70 0.649
6. Caudal peduncle depth 79 0.08-0.11 0.096
7. Head length 79 ~=—-0.25-0.31 0.287
8. Dorsal-fin base 79 ~=—-0..12-0.16 0.138
9. Dorsal-fin length 72 ~=0.24—0.33 0.283
10. Anal-fin base 78 0.42-0.53 0.486
11. Pectoral-fin length 79 = 0.17-0.28 0.250
12. Pelvic-fin length
13. Snout length 79 ~=0.18-0.26 0.223
14. Eye diameter 79 ~=0.33-0.44 0.393
15. Interorbital width 79 = -0.12-0.21 0.176
16. Upper jaw length 79 ~=0.41-0.57 0.486
Pristigaster whiteheadi
SD n Holotype Range Mean SD
— oy ‘S3o.4 22.0—86.0 56.12 —
0.041 95 0.56 0.42-0.58 0.534 0.031
0.014 95 043 0.40-0.48 0.425 0.017
O.0l2., O35 0:27 , 30.24-0.31,, 0272. >, 0.013
95 O59. “OAS-O.6Ok O57) . 0025
O;025: -95 0.67 0.54-0.71 0.657 0.024
0.008 95 0.09 0.07-0.12 0.098 0.007
DOIG" -95. 9 1029". O25-0:32-' 0.286 0:012
D009 95 euOil2s 0:12-0:17 ~ 0.145) 0012
0.020 94 0.26 0.24-0.32 0.289 0.018
0.026 95 0.44 0.42-0.50 0.467 0.016
UG. 3s 0.28 U.16-0°26° “0.230 ~“OOT
94 0.04 0.02-0.07 0.048 0.006
O02) “95 0.25 0.21-0.29 0.247 0.015
0.022 95 0.40 0.34—-0.45 0.399 0.023
0.016) +956» 0.20. -018-0.25.... 0:205.., 0.013
0.033: 95, , O49 ©.43-0:53 0:479. . 0.019
242
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Le:
Bigeile
Dorsal fin triangular when extended,
pointed dorsally; origin located on anterior
half of SL, and anterior to vertical through
anal-fin origin. Dorsal-fin rays 11+(12)—13,
third ray (unbranched) longest, twice as
long as fin-base. Pectoral-fin origin located
at middepth of body, slightly dorsal to ven-
tral margin of gill cover and slightly ante-
rior to its posterior margin. Pectoral-fin
length shorter than HL, its posterior tip,
when adpressed to body, reaching beyond
vertical through dorsal-fin origin. Pectoral-
fin rays 1+(10)—11. Axillary scale present
on region dorsal to pectoral-fin base, ex-
tending for approximately 20% of fin. Pel-
vic fins minute, length spanning approxi-
mately the space of three consecutive ab-
dominal scutes, with origin located nearly
at ventral margin of body, at vertical
through posterior tip of pectoral fin. Pelvic-
fin rays i+3+i. Anal fin long and low, its
origin slightly posterior to vertical through
posterior end of dorsal-fin base. Anal-fin
progressively shorter posteriorly. Tips of
last anal-fin rays reaching base of inferior
caudal-fin basal fulcra. Caudal fin deeply
Pristigaster whiteheadi, new species, holotype, MZUSP 52963, 83.4 mm SL.
forked, lower lobe slightly longer than up-
per. Caudal-fin rays 10/9. Vertebrae 43 or
44.
Scales large (approximately same size as
pupil) and cycloid, covering all of body,
and easily detachable, rarely preserved in
their entirety. Vertical scales rows 36-39.
Horizontal rows 18—20. Scale covering ex-
tending to midlength of middle caudal-fin
rays. Entire anal-fin base rimmed by row of
small scales (about half as large as remain-
ing body scales).
Pigmentation in alcohol.—Dorsum,
snout and upper part of sides brown. Re-
mainder of body silvery due to heavy de-
posits of guanine (which tend to disappear
after extended preservation). Narrow con-
centrations of dark melanophores on upper
and lower lips, and sometimes anterior por-
tion of chin. Region corresponding to neu-
rocranium dark, due to brain pigment visi-
ble through translucent skull bones. Scat-
tered dark chromatophores along entire dor-
sum and upper sides, more concentrated
along dorsal-fin base and dorsal part of cau-
dal peduncle. A middorsal dark spot is
sometimes present anterior to dorsal fin (as
VOLUME 113, NUMBER 1
avec
7 ee
Fig. 2.
/
ave
B
Predorsal bones and corresponding neural spines in lateral view, anterior to left. A—Pristigaster
cayana, MZUSP 30338; B—P. whiteheadi, n. sp., paratype, INPA 8555; Abbreviations: pb—predorsal bone,
ns—neural spine, avc—main axis of vertebral column. Scale bars = | mm.
in holotype). Myosepta visible but not out-
lined by dark pigmentation. Fins hyaline,
except for faint rows of melanophores
along outer rays of caudal fin and a few
scattered isolated melanophores on dorsal
fin.
Etymology.—The name of this species is
a posthumous tribute to Peter J. P. White-
head, who contributed more than any other
individual to the knowledge of clupeo-
morph fishes, also for first suggesting that
there might be two different species in Pris-
tigaster.
Distribution.—Brazil and Ecuador, in the
following river basins: Rio Solim6es/Ama-
zonas, Rio Trombetas, Rio Branco, Rio Ja-
pura, Rio Negro, Rio Madeira, Rio Jurua,
Rio Araguari (State of Amapa, Brazil) and
Rio Napo.
Pristigaster cayana Cuvier, 1829
Figs. 3, 4
Pristigaster cayanus Cuvier, 1829: pl. 10,
fig. 3; 1829:321 (name only, American
seas); Valenciennes, 1847 (redescription
of holotype).
Pristigaster argenteus Schinz, 1822:300
(based on Cuvier’s figure; nomen obli-
tum).
Pristigaster lichtensteinii Jarocki, 1822:
322, fig. 3 (based on Cuvier’s figure, re-
produced; nomen oblitum).
Pristigaster triangularis Stark, 1828:408
(based on Cuvier’s figure; nomen obli-
tum).
Pristigaster martii Agassiz, in Spix & Ag-
assiz, 1829:55, pl. 24a (Amazon); no pel-
vic fins; post-dates cayanus by a month
or so (see Whitehead & Bauchot, 1985);
Amaral-Campos, 1941:187; Whitehead
é& Myers, 1971;-Le Bail et al.,. 1983.
Pristigaster americanus Guérin-Menéville,
1844:33, pl. 57, fig. 3 (Atlantic coast of
South America).
Pristigaster phaeton Valenciennes, 1847:
338 (Amazon); no pelvic fins (see White-
head & Bauchot, 1985).
Pristigaster cayana; Myers, 1956 (validity
of P. cayanus Cuvier as of 1829; em-
mendation of specific name to agree with
feminine gender of genus); Hildebrand,
1964 (synonymy of cayana, phaeton and
martii); Whitehead, 1964:428, fig. 108
(synopsis); 1967:100, 102 (types of cay-
ana and phaeton); 1973a:85 (triangularis
244
a nomen oblitum; synonymy); 1985:301
(in part, only specimens without pelvic
fins; synonymy, diagnosis; distribution;
habitat and biology); Whitehead & My-
ers, 1971:487 (validity of martii); White-
head & Bauchot, 1985:24 (in part, only
specimens without pelvic fins; types of
cayana, phaeton); Eschmeyer, 1998:347
(catalog and suggestion—not accepted
here—that name is not available from
Cuvier, 1829).
Material examined.—95_ specimens
(23.5-142.0 mm SL). Brazil. Amazonas:
MZUSP 31032 (1, 90.0), Lago do Prato,
Rio Negro, Anavilhanas; MZUSP 11389 (1,
142.0), Lago Puraquequara; MZUSP 52947
(1, 64.0), Lago Manacapuru; MZUSP 7023
(20, 52.5—-74.0, 2 of which cleared and
counterstained), Rio Madeira, 25 kilometers
below Nova Olinda; MZUSP 11405 (1,
37.5); Rio, Solimoes,. Ilha do -Xibeco;
MZUSP 9568 (1, 96.0), Manaus; MZUSP
18696 (1, 92.0), Rio Solimdes, Lago Jan-
auaca; MZUSP 52943 (1, 23.5), Rio Negro,
1°58: 167S3261- 115/A22W:MZUSP52944.(6,
30.0—36.0), Rio Icd, 3.:O3.45,09,
68°04'26"W; MZUSP 52945 (1, 34.0), Rio
SolimGes, 2°40'15”S, 66°39'14"W; MZUSP
52946 (1, 62.2), Rio Japura, Parana do Ja-
pura, 3°08'20"S, 64°46'52”W; MZUSP 1388
(1-11 1.0),and 11390<(1,;36.0); Rio: Jurua:
BMNH 1925.10.28.5 (1, 88.9), Rio Soli-
moes, Manacapuru. Roraima: MZUSP
30334 (1, 137.0), Rio Branco, below Xe-
ruini; MZUSP 30335 (1, 112.0) and 30337
(1, 103.0), Rio Branco, Marara; MZUSP
30339 (2, 100.0 and 110.0), Rio Branco,
Lago do Maguari; MZUSP 30340 (6, 68.3—
82.5), Rio Branco, Xeruini; MZUSP 52942
(1,. 105-0):,. Rio 26Branceo,. 1 7:6.59"S,
61°50'52”W. Rondonia: MZUSP 30336 (1,
81.0) and 30338 (29, 33.0—66.0, 3 of which
cleared and counterstained), Rio Madeira,
Calama. Para: MZUSP 5560 (2, 72.0 and
73.5), Lago Ururié, Oriximina; MZUSP
5668 (1, 82.5), Lago Puru, Oriximina;
MZUSP 8280 (1, 97.0), Rio Trombetas,
Oriximina; MZUSP 5689 (1, 72.0), Rio
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Trombetas, mouth of Lago Paru. Mato
Grosso: MZUSP 17030 (1, 29.0), Rio Ar-
aguaia, Santa Terezinha; MZUSP18627 (2,
116.0 and 126.0), Rio Araguaia, Lago
Dumba. PERU. Loreto: FMNH 71264 (1,
100.0), mouth of Rio Tigre, 80 miles SW
of Iquitos; MZUSP 15217 (1, 83.0), Cocha
Aguajal, Rio Amazonas, Iquitos: MZUSP
15216 (2, 83.0 and 88.0), Rio Marafion,
Nauta; Pucallpa: MZUSP 18557 (1, 47.0),
Rio Ucayali.
Diagnosis.—See diagnosis of P. white-
headi.
Description.—Meristics and morphomet-
rics are presented in Tables 1 and 2. Body
form and disposition, shape and arrange-
ment of abdominal scutes (30-35 in num-
ber) as in P. whiteheadi.
Shape, size and position of snout, eye,
pupil, adipose eyelid, mouth, maxilla and
also shape, arrangement and number of
teeth as in P. whiteheadi. Number of teeth
also increasing with growth. Premaxillary
teeth 9 (in specimens <25 mm SL) to 23
in larger specimens. Maxillary teeth 18 (in
23.5 mm SL specimen) to 80 Gin 105 mm
SL specimen). Dentary teeth 7 (at 54 mm
SL) to 14 in larger specimens.
Shape of gill cover, gill membrane and
shape and disposition of gill rakers and po-
sition of dorsal limit of gill opening as in
P. whiteheadi. Number of gill rakers also
increasing with growth, ranging from 21 (at
45 mm SL) to 25 (at 110 mm SL) on lower
part of first branchial arch and from 9—12
on upper part.
Pelvic fins absent. Position, shape and
size of all other fins and axillary scale iden-
tical to those of P. whiteheadi. Dorsal-fin
rays 111+12—13. Pectoral-fin rays i+10—12.
10/9. Tips of dorsalmost two branched up-
per lobe caudal-fin rays elongated into fil-
ament twice as long as first principal ray
(unbranched) in a specimen 100 mm SL.
Tips of eighth and ninth lower lobe caudal-
fin rays also prolonged into a shorter lower
lobe filament about one-third as long as
tenth (unbranched) ray in same specimen.
VOLUME 113, NUMBER 1
25
20 A A At
ds,
A A A
= A A AA
2 AA A
¢ 15 A oOo
* Oo Oo
& Oo
2
Qa
=
3 10 Oo
oOo
=
e)
2
=)
i¢) ca = all a RJ a - er — ee ee mee ee
0 20 40 60
Fig. 3.
Upper and lower caudal-fin filaments fre-
quently broken in preserved specimens.
Vertebrae 43-44.
Size, shape and distribution of cycloid
scales on body and fins as in P. whiteheadi.
Vertical scale rows 40—47. Horizontal rows
21-26.
Pigmentation in alcohol.—As in P.
whiteheadi.
Distribution.—Nearly coincident with
that of P. whiteheadi, with which it is sym-
patric in most localities of the Amazon Ba-
sin, but extending further south into Rio
Araguaia, State of Mato Grosso, Brazil.
The locality associated with the holotype
of the species is reported as Cayenne,
French Guiana (Whitehead 1967). This in-
formation is not provided in the original
accounts on the species by Cuvier, which
State simply “‘seas of America’’. Other re-
ports on the type-locality have inferred it
from the species name (e.g., Myers 1956).
Representatives of Pristigaster have never
been collected again in French Guiana, in
any of the surveys of the area (e.g., Le Bail
245
o o
o o
Oo o
A oo Oo
ao oo Oo
Oo
Oo Oo
o 8a 5
o
O Other samples of P. cayana
4 Rio Madeira sample
|
|
80 100 120 140 160)
Standard lenght - mm
Size-dependent variation of the number of premaxillary teeth in P. cayana.
et al. 1983, Planquette et al. 1996, also P.
Keith, pers. comm.). The genus has also
not been reported from surrounding areas,
like Guyana, Suriname or the Orinoco ba-
sin. We strongly suspect that the Cayenne
locality is erroneous, and may simply re-
flect a port of shipment or an intermediate
post en route between South America and
France, in which the material studied by
Cuvier may have remained temporarily.
Remarks.—The sample from Rio Madei-
ra differs from remaining ones in number
of premaxillary teeth (Fig. 3), but no other
meristic or morphometric difference was
found between that population and others
throughout the range of the species. In the
absence of additional significant differences
we prefer to consider the higher number of
premaxillary teeth in the Rio Madeira sam-
ple as populational variation.
Notes on the name P. cayana.—The his-
tory of the names associated with the spe-
cies referred to here as Pristigaster cayana
is rather complicated. The first reference to
the species was done in Cuvier (1816), as
246
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Pristigaster cayana, MZUSP 30340; caudal fin in lateral view, showing filaments. Scale bar = 1 mm.
an illustration associated with the genus
name only. In 1829, Cuvier proposed the
name cayanus for the species figured in
1816, in what is clearly a valid species de-
scription as was shown by Myers (1956).
The species name therefore must date from
1829, and it precedes the validly described
P. martii by a month or so (Whitehead
1985:301). Various other names (argen-
teus, lichtensteinii and triangularis) were
proposed between 1816 and 1829, all on
the basis of the same illustration in Cuvier
(1816). We consider all of these names to
qualify as nomina oblita according to the
International Code of Zoological Nomen-
clature (ICZN, 1985, Art. 79c). They have
never been used subsequently as the valid
names for the species, and the junior syn-
onym, cayanus, has certainly been used in
more than 10 publications by more than
five authors in the intervening period (e.g.,
Valenciennes 1847, Giinther 1868, Jordan
& Evermann 1896, Norman 1923, Ihering
1930, Myers 1956, Whitehead 1967, 1973,
1985; Whitehead & Bauchot 1985).
Discussion
As demonstrated in this paper, in spite of
the overall similarity in body shape and pig-
mentation, Pristigaster cayana and P. whi-
teheadi are clearly distinct and diagnosable
through several morphological features.
When suggesting the recognition of a sec-
ond species of the genus, Whitehead (1985)
and Whitehead & Bauchot (1985) based
their observations on just a few specimens
and did not notice some of the striking dif-
ferences we found. In the diagnostic fea-
tures of P. cayana, Whitehead (1985) de-
scribed the pelvic fins as usually absent, in-
dicating that presence or absence could
vary within the species. At the same time,
however, the author considered that one
Species without pelvic fins and high gill
raker counts (22—24, usually 23) could pos-
sibly be different from a species bearing
pelvic fins with lower gill raker counts (19—
20, usually 20). Results of our study reveal
that presence of pelvic fins is indeed diag-
nostic. They exist only in P. whiteheadi and
VOLUME 113, NUMBER 1
are consistently present from the smallest to
the largest individual of both sexes. Gill
raker counts overlap to a certain extent, but
there is a significant mean difference (see
Table 1) between the two species. Other dif-
ferences pointed out in the diagnosis of P.
whiteheadi leave little doubt that there are
indeed two separate species in Pristigaster.
None of the various other names applied to
P. cayana (see synonymy above) could
possibly have been based on the new spe-
cies. They either were based on Cuvier’s
1816 illustration or clearly mentioned the
absence of pelvic fins in specimens exam-
ined.
The two Pristigaster species share the
highly peculiar expanded morphology of
the abdomen, to a degree which distinguish-
es them from all other recent clupeiforms.
Not only the depth, but also the shape of
the abdominal expansion (abruptly emerg-
ing nearly vertically from gular region) are
obviously apomorphic conditions not seen
elsewhere in other recent clupeomorphs,
and strongly suggest that Pristigaster is
monophyletic. These characteristics are as-
sociated with a host of internal-anatomical
modifications not yet studied in detail, and
which will be the subject of a forthcoming
paper.
Sexing of most of the specimens of both
species examined did not reveal any obvi-
ous sexual dimorphism. It also tested the
validity of our interspecific diagnostic char-
acters by showing that the differences were
not simply due to sexual dimorphism.
Acknowledgments
We thank Efrem Ferreira and Paulo Petry
(INPA), Tony Gill and Oliver Crimmens
(BMNH), and Barry Chernoff and Mary-
Anne Rogers (FMNH), for the loan of ma-
terial under their care. We are also grateful
to Philippe Keith for information regarding
collections in French Guyana. The manu-
Script benefited from reviews by Fabio Di
Dario, Lance Grande, Thomas Munroe and
an anonymous reviewer. Research funding
247
for both authors is provided by the Conse-
Iho Nacional de Desenvolvimento Cientifi-
co e Tecnolégico (CNPq; individual grants)
Projeto PRONEX, and Fundagao de Am-
paro a Pesquisa do Estado de Sao Paulo
(FAPESP).
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):249-—263. 2000.
A new species of Apogon (Perciformes: Apogonidae) from the Saya
de Malha Bank, Indian Ocean, with redescriptions of Apogon regani
Whitley, 1951, A. gardineri Regan, 1908, and A. heraldi
(Herre, 1943).
Thomas H. Fraser
W. Dexter Bender and Associates, Inc., 2052 Virginia Avenue, Fort Myers, Florida 33901, U.S.A.
Abstract.—A new species of fish, Apogon quartus, known only from the
Mascarene Plateau, is described. It is related to Apogon poecilopterus Cuvier
in Cuvier and Valenciennes, 1828, Apogon carinatus Cuvier in Cuvier & Va-
lenciennes, 1828, and Apogon queketti Gilchrist, 1903, recently treated by Gon
(1996) as members of the subgenus Jaydia Smith, 1961. Apogon quartus can
be distinguished from these three species by having a single predorsal scale,
18 total (13 well developed) gill rakers on the first arch, and a single dark spot,
perhaps appearing as an ocellus in life, on the body just behind the opercle
flap. Six additional nominal species are added to the 10 valid (19 nominal)
species treated by Gon in Jaydia. Three of these names, Apogon argyrogaster
Weber, 1909, formerly in Siphamia, Apogon melanopus Weber, 1911, and Apo-
gon fuscomaculatus Allen & Morrison, 1996, are valid species. The holotype
of Apogon heraldi (Herre, 1943) redescribed here, is a synonym of A. poeci-
lopterus. Apogon fuscovatus Allen, 1985 was determined to be a synonym of
A. melanopus by Allen & Morrison (1996) The other name, Apogon tchefouen-
sis Fang, 1942, may be synonymous with one of the species Gon treated but
is not placed with certainty. Character overlaps between the subgenus Jaydia
and other Apogon subgenera, particularly the largest subgenus Ostorhinchus
Lacépeéde, 1802, are briefly examined. Virtually all derived characters of Jaydia
grade into Ostorhinchus. Two rare species Apogon regani Whitley, 1951 and
Apogon gardineri Regan, 1908 only known from the Mascarene Plateau are
redescribed from new material. Both belong with the Apogon nigripinnis Cu-
vier in Cuvier and Valenciennes, 1828, complex of species.
The Russian vessel Vityaz made trawl
stations along the Saya de Malha Bank dur-
ing 1989 collecting fishes. Two of these
collections contained a new species and a
specimen of Apogon regani Whitley, 1951.
The new species was first identified as an
Apogonichthys Bleeker, 1854c, based on the
mostly smooth preopercle edge and the
slightly rounded caudal fin. An examination
of internal and external characteristics led
to the conclusion that this specimen is an
Apogon Lacépéde, 1802. Some characters
are held in common with species in the sub-
genus Jaydia recently treated by Gon
(1996). The ten species of Jaydia have the
fourth dorsal spine longer than any other
dorsal spine, variable serrations on pre-
opercular edges, rounded or truncated cau-
dal fins, and some have bioluminescent or-
gans. Gon and Allen (1998) included an-
other new species, Apogon photogaster in
Jaydia. A review of additional nominal spe-
cies with possible relationships to the new
species yielded six names, Apogon argyr-
ogaster Weber, 1909, (previously in Si-
phamia Weber, 1909, see Lachner 1953, p.
250
416), Apogon melanopus Weber, 1911,
Apogon tchefouensis Fang, 1942, Apogon
heraldi (Herre, 1943), Apogon fuscovatus
Allen, 1985 and Apogon fuscomaculatus
Allen & Morrison, 1996.
Specimens of all the above species have
been most frequently taken by trawls, often
in deeper waters. Fishes in Jaydia have
body shapes ranging from elongate more
slender forms, Apogon truncatus Bleeker,
1854e, to A. melanopus a deeper body form
with an emarginate caudal fin typical of
most Apogon. Smith (1961) based Jaydia
on Apogon ellioti Day, 1875, now recog-
nized as a synonym of A. truncatus a slen-
der species with a rounded caudal fin and
having bioluminescent activity.
In the process of trying to locate more
material of the new species, specimens of
two deep-dwelling species from the Naza-
reth Bank near Cargados Carajos Shoals
(St. Brandon) described by Regan (1908)
were found. Randall and van Egmond
(1994) reported Apogon punctatus Regan,
1908, (=Apogon regani Whitley, 1951) tak-
en in a trawl from the Seychelles. No ma-
terial of Apogon gardineri Regan, 1908, has
been reported since the original description,
but deep dives by J. E. Randall during 1979
in Mauritius yielded two specimens. It is
likely that more material will be only slow-
ly forth coming. Both of these species are
regarded as endemics of the Mascarene Pla-
teau.
Methods
Methods of taking and recording meristic
data and measurements are given in Fraser
& Lachner (1985). All measurements are in
millimeters to the nearest 0.1 mm. All pro-
portions are based on standard length and
all material is reported by standard length
rounded to the nearest millimeter, except
for the primary type material. All x-ray
photographs are in data files maintained by
the author. The acronyms used in the lists
of materials to designate institutions and
collections cited, follow general usage giv-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
en in Leviton et al. (1985) and Eschmeyer
(1998).
Apogon quartus, new species
Figs. [dé 2
Material examined.—Holotype: USNM
307688; (49.8); Indian Ocean, Saya de Mal-
ha Bank, 11°05’00"S, 62°02’00"E; Vityaz
Cr. 17; Sta 2808; 8 Jan 1989; 58—61 m.; X-
ray.
Comparative material.—Amia albomar-
ginata Holotype: USNM 68402; (83.6);
Philippines, Cavite; X-ray. Apogon arafur-
ae Holotype: BMNH 1879.5.11.141; (90.0);
Arafura Sea. Apogon argyrogaster Syn-
types: ZMA 101075; (34.9—47.5); New
Guinea, west coast; Siboga sta 164; 32 m.
BPBM 32628; (15.5—48.3); New Guinea,
Nagada Harbor; 18 Nov 1987; 30 m; X-ray.
Apogon bilaciniatus Lectotype: ZMA
101280; (46.2); Indonesia, Lombok. Apo-
gon ellioti Paralectotype?; ZSI 1905; (77.7,
96.0 mm TL); India, Madras; X-ray. Para-
lectotype?; AMS B.8226; (77.3, 98.1 mm
TL); India, Madras; X-ray. Apogon fusco-
vatus Holotype: WAM P14397; (94); Aus-
tralia, Darwin; 4 Sep 1965. Paratypes:
WAM P14516—-17; (83-85); Australia, N.
of Darwin; 9 Sep 1965. WAM P 28316-
001; (66); Australia, Darwin; 10 Sep 1965.
Apogon glaga Syntype: RMNH 5614 (71.9,
~93 mm TL); Indonesia. Apogon hungi
Neotype: USNM 340009; (76.4); Mozam-
bique Channel. Jaydia hungi Holotype:
MNHN 1965-711 (94.3); Egypt, Gulf of
Suez. Apogon lineatus Lectotype: RMNH
70a; (62.4); Japan. USNM 71240; (64.6);
Japan, Shimizu Suruga; Albatross; 1906;
female. USNM 32586: (66.3): China:
35°55'49"N,. 120°2129°E; 25 Jule 1993 -ie-
male. Amia melas [=Apogonichthys melan-
opterus| Holotype: ANSP 47491; (~35.2
head distorted); Philippine Islands. Apogon
melanopus WAM P. 14963; (100); Austra-
lia, Darwin; 4 Sep 1965. Apogon modestus
Holotype: RMNH 5579; (61.9); Indonesia,
Java. Apogon nigricans Syntype: ZSI 1872;
(55.0, 71+ TL) India, Madras; X-ray. Apo-
VOLUME 113, NUMBER 1
Hire...
gon novaeguineae Holotype: MNHN 8695;
(56.5); New Guinea. Apogon photogaster
Paratype: USNM 348214; (43.3); Papua
New Guinea, Madang Lagoon; 21 Oct
1996; 18—23 m; X-ray. Apogon poecilop-
terus Holotype: RMNH 214; (79.0); Indo-
nesia, Java. Apogon queketti Syntypes:
SAM 11657; (75.1); South Africa, Natal;
X-ray. SAM 11658; 5(44.1-—77.4); same
data; X-ray. Jaydia smithi Paratypes: ZMH
5034; 5(12.7—46.3); Amia striata Holotype:
USNM 68403; (67.2); Philippines, Luzon;
female; X-ray. Paratypes: USNM 93410;
11(39.2—66.2); same data; X-ray. Somalia,
Gulf of Aden; X-ray. Apogon striatodes
Holotype: USNM 213408; (55.5); Indian
Ocean, Thailand. Paratypes: China, Hong
Kong: CAS 160877; (57); CAS 161015;
(55). Philippines, Luzon: CAS 85669; (45);
Corregidor I.: CAS 32723 (46). Thailand,
Gulf of Thailand: CAS 79648; (36-37);
CAS 79652; (44); (CAS 82205; 8(18—-59);
CAS 85676; (26-30); CAS 85666; 3(28—
34); CAS 85667; (19); CAS 82202; (17-
47); CAS 82208; (54). Apogon tchefouensis
Paratypes: MNHN 1941-148; (45.7); China,
Shandong Prov., Tché-Fou, Ho Ting Chieh;
X-ray. MNHN 1941-149; (35.0); same data;
X-ray. Apogon truncatus Holotype: RMNH
5582; (55.4); Indonesia, Batavia (=Jakarta).
The holotype of Apogon quartus from the Saya de Malha Bank, Indian Ocean, 49.8 mm SL.
Apogon hoevenii Syntype: RMNH 5581;
(40:2; 50m“ TE; one ‘of: 6"'spec: imisize
range, 24 spec. 25—49 mm SL); Indonesia,
Ambon. USNM 261058; 8(35—39); Philip-
pines, Siquoijor I. Apogon sp. USNM
349199; (48.8); Philippines, Palawan,
Puerto Princesa City.
Diagnosis.—A species of Apogon in the
subgenus Jaydia with 15 pectoral rays, gen-
erally smooth to crenulate preopercle edge,
one predorsal scale, 18 total (13 well de-
veloped) gill rakers on the first arch, fourth
dorsal spine the longest, and a single dark
spot, perhaps appearing as an ocellus in life,
on the body just behind the opercle flap.
Description.—For general body shape
see Fig. 1. Proportions (as percentage of
standard length): greatest body depth 33.5;
head length 42.2; eye diameter 11.4; snout
length 9.2; bony interorbital width 10.2; up-
per-jaw length 20.5; caudal-peduncle depth
14.8; caudal-peduncle length 21.5; first dor-
sal-spine length 3.4; second dorsal-spine
length 9.0; third dorsal-spine length 15.7;
fourth dorsal-spine length 17.9; spine in
second dorsal fin 14.8; first-anal-spine
length 2.4; second-anal-spine length 11.4;
pectoral-fin length 21.2; pelvic-fin length
22.
Dorsal fin VII-I,9; anal fin 1,8; pectoral
252
fin 15-15; pelvic fin I,5; principal caudal
rays 9 + 8; number of simple pored lateral-
line scales unknown, series extending from
posterior edge of posttemporal to caudal
fin; transverse scale rows above lateral line
2: transverse scale rows below lateral line
~4; median predorsal scales 1, the lateralis
system extending well onto the nape as a
raised semi-translucent fleshly area of about
seven striations or rows with many small
papillae; number of circumpeduncular scale
rows unknown; total gill rakers 18, well de-
veloped 13, 342 upper arch,. 11--2 lower
arch.
Villiform teeth in wide band on premax-
illa; wide band grading to two rows on den-
tary; one to two rows on palatine and one
to three rows on vomer; none on ectopter-
ygoid, endopterygoid or basihyal.
Vertebrae 10 + 14. Five free hypurals,
one pair of slender uroneurals, three epur-
als, a free parhypural. Three supraneurals,
two supernumerary spines on first dorsal
pterygiophore. Basisphenoid present. Su-
pramaxilla absent. Posttemporal smooth on
posterior margin. Preopercular ridge
smooth, posterior vertical edge smooth,
ventral horizontal edge slightly crenulate
with one or two spinelets at angle. Infra-
orbital with irregular edges, but without ser-
rations. Scales ctenoid. Caudal fin slightly
rounded. Nose and pre-interorbital area
semi-translucent and slightly bulbous. No
indication of bean-like bioluminescent or-
gans associated with intestine near anus.
Life colors.—Unknown.
Preserved color pattern.—In 70% ethyl
alcohol: peritoneum pale, stomach and in-
testine pale. Dark mark from ventral margin
of eye ending near posterior edge of pre-
maxilla, two dark lines from posterior mar-
gin of eye, upper to posttemporal, lower
onto opercle ending in a ventrally directed,
short mark; dark spot ocellus-like on side
just behind opercle flap, with about five
smaller spots positioned anteriorly and ven-
trally close to large spot; other melano-
phores on pectoral-fin base and abdomen,
scattered diffuse spots on caudal peduncle
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
behind second dorsal and above anal fin;
scales above the lateral-line scales outlined
with melanophores; lower third of snout
dusky between upper jaw and eye; pelvic
fins with spine and distal third of rays dark
to tips; anal fin with dark band on spines
and first three fin-rays to tip, fourth fin-ray
mostly dark, distal third of next two fin-rays
dark, last two fin-rays pale from base to
tips; caudal fin with tips of rays dark; sec-
ond dorsal-fin spine and first four fin-rays
dark to tips, remaining fin-rays pale; first
dorsal fin with the anterior four spines and
membranes dark to their tips, remaining
spines and membranes pale.
Distribution.—Known from the Indian
Ocean on the Saya de Malha Bank (Fig. 2).
Etymology.—The Latin word for fourth,
quartus in reference to the longest dorsal
spine.
Remarks.—The fourth dorsal spine is the
longest dorsal spine in this specimen. With
additional characters including a high num-
ber of pectoral-fin rays, rounded caudal fin
and preserved color pattern, this species can
be placed within the broad limits of sub-
genus Jaydia revised by Gon (1996). Apo-
gon quartus is the first species of this sub-
genus to be found on the extensive isolated
shallow banks the Central Indian Ocean.
Most of the species treated by Gon (1996)
have a continental distribution pattern. Only
a few specimens of Apogon hungi Four-
manior & Nhu-Nhung, 1965, and Apogon
smithi (Kotthaus, 1970) have been reported
from insular localities (Gon, 1996).
Apogon quartus appears to be most
closely related to Apogon poecilopterus Cu-
vier in Cuvier & Valenciennes, 1828, Apo-
gon carinatus Cuvier in Cuvier & Valen-
ciennes, 1828, and Apogon queketti Gil-
christ, 1903. The new species differs from
A. queketti and A. carinatus by not having
an ocellus in either dorsal fin. A. poecilop-
terus has more predorsal scales (4—5) and a
darkish gill chamber. The ocellus-like spot
on the body is unique to A. quartus among
these species.
Apogon argyrogaster is unique among
VOLUME 113, NUMBER 1
40° 60°
Arabian Sea
Lakshadweep Is.
Africa
Mauritius
Madagasgar
Maldive ‘3
Is. 2
80° 100°
Re Bay of Bengal age
“3 Andaman
vt ay
Sri acess ‘s
Lanka S.
Jawa Sea
26
Cocos (Keeling) ; .
Is. . Christmas
~ Rodrigues
* Apogon quartus
@ Apogon regani
@ Apogon gardineri
40° 60°
Fig... 2.
species of Apogon with its large ventral or-
gans (probably bioluminescent) extending
on each side of the body from the breast
area forward into the lower part of the
mouth and backward to past the origin of
the anal fin. Gon & Allen (1998) recently
described Apogon photogaster with an ex-
ternally similar bioluminescent system to A.
argyrogaster but made no comparisons.
Their name appears to be a synonym of A.
argyrogaster. They placed A. photogaster
in Jaydia. Apogon argyrogaster has faint
bars on the side of its body, two predorsal
scales, 8—9 well developed gill rakers in ad-
dition to the luminous organs. All are char-
acters differing from A. guartus.
Apogon melanopus, with nine anal rays
80° 100° 120°
Distribution of collection sites for Apogon quartus, Apogon gardineri, and Apogon regani.
and an emarginate caudal fin unlike all oth-
er Jaydia species, is a member of Jaydia
with the fourth dorsal spine the longest and
16-17 pectoral-fin rays. This species was
described by Allen (1985) as A. fuscovatus.
Allen & Morrison (1996) later synony-
mized Allen’s species when describing an-
other new species, A. fuscomaculatus,
which they compared with Apogon striatus
(Smith & Radcliffe in Radcliffe 1912).
Apogon melanopus and A. fuscomaculatus
differ in having higher well-developed gill
rakers (15—16) and lower well-developed
gill rakers (8—9) respectively and general
color pattern (no ocellus-like spot in either
species) from A. quartus.
Paratypes of A. tchefouensis from the
254
East China Sea examined by me have no
discernable color patterns because of the
strong brown stain on both specimens. Fang
(1942) described the color in alcohol as
5 uniformément brun-noiratre.”’ Fang
(1942) described the preopercle as “Bord
libre du préopercule nettement denticulé
avec des dents assez fortes surtout a son
angle, mais plus fines a la partie supérieure
du bord postérieur.’? The condition of the
preopercle with a smooth ridge and prom-
inent serrations, stronger at the angle and
finer ones on the posterior edge and ventral
edge, suggest a different group, probably
type B (but serrations more like type A on
edges) of Gon’s categories than with the
group (type C) containing A. quartus. The
posttemporal is serrated. Total gill raker and
rudiment count on the first gill arch is 18,
well-developed rakers 14 (3+2 on upper
arch, 12+1 on lower arch) of the larger
paratype. The smaller paratype is in poor
shape and has a badly damaged head. The
pectoral ray count was 15 on one side of
both paratypes and 14 for the other side for
the larger paratype. Palatine teeth are in one
to two short rows for the larger paratype.
Palatine teeth of the lectotype of Apogon
lineatus Temminck & Schlegel, 1842, are
in 4 short rows like the holotype of Apogon
striatodes Gon, 1996, (3—4 short rows), and
the holotype A. striatus (3—4 short rows)
but not Apogon novaeguinae Valenciennes,
1832 with one row. Additional information
about the color of the peritoneum and in-
testine from the holotype and paratypes
may help determine whether A. tchefouen-
sis 1S a junior synonym of either A. lineatus
or A. striatus or the senior synonym of A.
striatodes. The known distribution of the
two species recognized by Gon (1996) in
the East China Sea region do not appear to
overlap, but he suggests the possibility for
Taiwan. If distributional patterns are accu-
rate, then the larger paratype of A. tche-
fouensis should be identified as A. lineatus.
What ever the eventual status of A. tche-
fouensis, the paratypes can be distinguished
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
from A. quartus by having serrated preoper-
cle and posttemporal.
Fowler & Bean (1930) compared Apo-
gonichthys melanopterus, their replacement
name for Amia melas Fowler, 1918, with
Apogon carinatus. Fowler (1918) first re-
lated his new species to Apogon nigricans
Day, 1875, a nominal species related to, if
not the same as Apogon melas Bleeker,
1848, with a forked caudal fin, a trace of a
dark spot on the midbase of the soft dorsal
fin and darkish pelvic and vertical fins. In
his original description, Fowler noted that
the holotype of Amia melas has the fourth
dorsal spine longer than the third. My ob-
servation of the holotype agrees with Fowl-
er’s statement. However, the third spine is
noticeably stronger (thicker) than the fourth
spine unlike other species of Jaydia. Apo-
gon melanopterus 1s consistent in some oth-
er characters with Jaydia: gill-raker count
(3+1-11+3), some serrations on the pre-
opercle edge and a rounded? (damaged)
caudal fin, but has 14 pectoral-fin rays ver-
sus the common 15-17, rarely 14 or 18 pec-
toral-fin rays of Jaydia. This species differs
from A. quartus in preserved color pattern
(no ocellus-like spot; dorsal, pelvic, anal
and caudal fins all blackish), predorsal
scales (3), strong third dorsal spine and 14
pectoral rays. I cannot with certainty place
A. melanopterus within Jaydia as treated by
Gon.
Relationships.—The basis for the pro-
posed monophyly of Jaydia will need to be
re-examined because some of Gon’s (1996)
diagnostic features are present in other spe-
cies of Apogon. For example, Apogon hoev-
enii Bleeker, 1854d, a more ‘typical’ Apo-
gon with an emarginate caudal fin, serrated
preopercular edge and a low pectoral ray
count (12), has the fourth dorsal spine var-
iably as its longest. The holotype of A. me-
lanopterus has the fourth dorsal spine as the
longest and a robust thicker third dorsal
spine. One undescribed species and a relat-
ed species, Apogon moluccensis Valenci-
ennes, 1832, have the fourth dorsal spine as
the longest in nearly all adults but not in all
VOLUME 113, NUMBER 1
small individuals. The third spine in these
two species is about as strong as the fourth
spine, and both have forked caudal fins. The
undescribed species has a darkish roof of
the mouth and gill arches reminiscent of A.
poecilopterus. An unidentified species of
Apogon, either new or a juvenile of a spe-
cies in the Apogon diversus (Smith & Rad-
cliffe in Radcliffe, 1912) group, has strong
serrations on the preopercular ridge and
edges, serrated infraorbitals, a strongly fork
caudal fin and fourth dorsal spine as the
longest spine. Characters present in A. me-
lanopus and the above species suggest that
an expanded review is needed. A solid basis
for the subgenus founded on the shape of
the caudal fin, the length of the fourth dor-
sal spine, strength of the third dorsal spine,
high pectoral-fin ray counts, supporting
structures for the dorsal fin and other char-
acters elucidated by Gon may grade with
the examination of more species. Perhaps
some of the other species of Apogon dis-
cussed here eventually may be shown to be
closer to this species complex than to other
Apogon. Those species treated by Gon that
have bioluminescent systems are more like-
ly to be monophyletic and may be worthy
of the recognition as the subgenus Jaydia.
There is, as yet, no clear resolution about
whether these two subgenera represent
monophyletic groupings within Apogon.
Status of Apogon heraldi
Fig. 3
Apogon heraldi (Herre, 1943) was de-
scribed in Mionurus, a misspelling of Mio-
norus Krefft, 1868, as having nine anal
rays. Herre compared it with two unrelated
nominal species with six first dorsal spines,
Apogon bombonensis (Herre, 1925) and
Apogon mydrus (Jordan & Seale, 1905).
There is enough information in Herre’s de-
scription to determine that this species
should be compared with those treated by
Gon (1996), but not enough to place it with
certainty. Herre did not give a figure of the
type. A redescription of the type, with a fig-
255
ure 1s needed to provide comparison with
the new species and with other related spe-
cigs,
Material examined.—Mionurus heraldi
Holotype: SU 38263 (104.5); Philippines,
Luzon I., Ragay Gulf, X-ray.
Comparative material.—See A. quartus.
Description.—For general body shape
see Fig. 3, Range of proportions (as per-
centage of standard length): greatest body
depth 39.4; head length 42.0; eye diameter
9.8; snout length 8.5; bony interorbital
width 9.6; upper-jaw length 20.1; caudal-
peduncle depth 16.7; caudal-peduncle
length 20.2; first dorsal-spine length 2.4;
second dorsal-spine length 7.1; third dorsal-
spine length 10.9; fourth dorsal-spine
length 14.1; spine in second dorsal fin 11.3;
first anal-spine length 2.8; second anal-
spine length 10.9; pectoral-fin length 25.4;
pelvic-fin length 26.7.
Dorsal fin VII-—I,9; anal fin II,8; pectoral
fin 16-16; pelvic fin I,5; principal caudal
rays 9 + 8; number of pored lateral-line
scales unknown; number of transverse scale
rows above lateral line unknown; number
of transverse scale rows below lateral line
unknown; median predorsal scales 4 or 5;
number of circumpeduncular scale rows un-
known. Gill rakers, upper arch 2 rudiments
1 well developed raker, lower arch | rudi-
ment, 10 well-developed rakers, 11 well de-
veloped and 14 total.
Villiform teeth in single row anteriorly,
slightly larger grading into wide band at
sides of premaxilla; three rows anteriorly
with one to two rows at sides of dentary,
interior row slightly larger from anterior to
side; one row on palatine; two rows on vo-
mer; none on ectopterygoid, endopterygoid
or basihyal.
Vertebrae 10 + 14. Five free hypurals,
one pair of slender uroneurals, three epur-
als, a free parhypural. Three supraneurals,
two supernumerary spines on first dorsal
pterygiophore. Basisphenoid present. Su-
pramaxilla absent. Posttemporal smooth on
posterior margin. Preopercular ridge
smooth, vertical margin smooth, horizontal
256
The holotype of Mionurus heraldi, SU 38263, 104.5 mm SL from the Philippine Islands, Luzon I., Ragay Gulf.
Fig. 3.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
margin with smooth undulations. Infraor-
bitals with crenulate to sharp edges. Scales
ctenoid, many missing from body. Caudal
fin slightly rounded. No indication of bean-
like bioluminescent organs associated with
intestine near anus.
Life colors.—Unknown.
Preserved color pattern.—In 70% ethyl
alcohol: peritoneum silvery; intestine and
stomach mostly pale with scattered mela-
nophores; second dorsal fin with dark stripe
near base, a narrow pale area above on first
few rays to about fifth ray, more distally the
fin-ray membranes dusky to dark with sec-
ond narrow pale area, then fin rays dusky
to tips; wide darkish stripe basally in anal
fin, the membranes pale distally; spinous
dorsal dusky distally, dark between the sec-
ond and fifth spines, pale basally; pelvic
and pectoral fins pale; gill chamber and gill
arches dusky with melanophores; caudal fin
membranes dusky with no indication of a
darker margin.
Remarks.—Apogon heraldi has the
fourth dorsal spine longest and can be
grouped with Gon’s type C preopercle pat-
tern of Jaydia. Herre (1943) described the
color of A. heraldi in alcohol as “‘... pale
brown, with a more or less evident vertical
dark-brown bar under each scale; there is a
blackish spot on the upper half of the spi-
nous dorsal, and a wide dark-brown cross-
band on the second dorsal and the anal,
which are otherwise clear; all but the basal
part of the caudal is dusky; the pectorals
and ventrals are clear’. Herre’s description
of the teeth as “The very small conical
teeth are in a single row in the lower jaw,”
and lack of comment about color in the oral
cavity would suggest that this specimen is
not A. poecilopterus. However, my re-ex-
amination of the holotype indicates that the
dentition and the color pattern in the gill
chamber are consistent with that of A. poe-
cilopterus. The holotype has 14 gill rakers
plus rudiments on the first gill arch, 2+1 on
the upper arch and 10+1 on the lower arch,
about 4 or 5 predorsal scales and 16 pec-
toral rays, all consistent with A. poecilop-
VOLUME 113, NUMBER 1
terus. Herre (1943) reported on material of
A. ellioti [=truncatus] and A. poecilopterus
from the Ragay Gulf and A. striatus from
Manila Bay in the same paper. Apogon her-
aldi is considered a synonym of A. poeci-
lopterus. The type is a ripe female.
Apogon gardineri Regan, 1908
Figs. 2 & 4
Material examined.—Holotype: BMNH
1908.3.23.93; 39.7 mm SL; Cargados Ca-
rajos; 55 m. Other material: BPBM 24775;
(41—42); Mauritius, off Flic en Flac; J. E.
Randall et al.; 7 April 1979; 57 m; color
photo; x-ray.
Comparative material.—Apogon nigri-
pinnis Syntype: MNHN 8694; 63.4 mm SL;
India, Pondicherry; Leschenault. Mozam-
bique: Delagoa Bay: RUSI 3149; (60-76);
x-ray. RUSI 3151; (67); 1938; x-ray. RUSI
3148; 4(42—84); x-ray. RUSI 3150; 4(68-—
74); x-ray. Palma: RUSI 3152; (69); 1 Aug
1951; x-ray. Aldabra I.: RUSI 3153; (49-—
53); 14 Nov 1954; x-ray. Apogon striatus
Syntypes: MNHN 1973-41; 3(16.8—26.2);
Madagascar, Nosy-Bé; x-ray. Apogon suezii
Hotetype: MNHN 5137; (50:6); Egypt,
Suez; Letourneux. Apogon taeniatus Syn-
types: MNHN 8693; (55.3-—57.4); Red Sea;
Ehrenberg. Kenya: RUSI 3106; (85); Mal-
medi; 2Oct 1952; x-ray. RUSI 3111; (94);
Shimoni; 1950; x-ray. RUSI 3107; (77-83);
Shimoni; Nov 1952; x-ray. RUSI 3112;
3(15—66); Shimoni; May 1951; x-ray. RUSI
3109; (76); Feb 1952; x-ray. Mozambique:
RUSI 3105; (79); Pinda reef; 1950; x-ray.
RUSI 3114; (49); Mocimboa de Praia; 3
Sep 1951; x-ray. RUSI 3110; (81); Inham-
bane; Sep 1948; x-ray. RUSI 3108; 3(59-—
82) Jun 1950; x-ray. Aldabra: RUSI 3113;
(55); Nov. 1954; x-ray. ANSP 63868; (86);
South Africa. Apogon thurstoni Holotype:
BMNH 1889.8.17.2; (55.5); India, Madras;
Day. Apogon timorensis Holotype: RMNH
5583; (60.8); Timor. QM 1.820; 3(23-31);
Australia, Darnley I. QM I. 8506; (28);
Australia, Queensland, Green I. Apogon-
ichthyoides fraxineus Holotype: RUSI 356;
pmo
(57.7); Mozambique, Pinda Reef; 3 Sep
1956; x-ray. Paratypes (all x-rayed): Moz-
ambique: RUSI 746; 8(42—65); Pinda Reef;
2 Sep 1956..RUSI 751; (67); Pinda Reef;
Sep 1956. RUSI 762; (37-69); Pinda Reef;
23 Sep 1956. RUSI 758; (48-55); Delagoa
Bay. RUSI 7409; (33); Ibo I.; Aug 1951.
Zanzibar: RUSI 757; (41); 20 Sep 1952.
RUSI 754; (54); 9 Sep 1952. RUSI 748;
3(52-57); 4 Sep 1952. RUSI 759; (23-24);
Seychelles, Mahe I.; Sep 1954.
Diagnosis.—A species of the subgenus
Ostorhinchus with two saddles on the body
and one on the caudal peduncle, no cheek
mark, a pale stomach, intestine and perito-
neum, 14 pectoral-fin rays, 2 predorsal
scales and 14—15 total gillrakers.
Description.—For general body shape
see Fig. 4. Range of proportions (as per-
centages of standard lengths), holotype in
parentheses: greatest body depth 38-39
(37.3); head length 42 (42.3); eye diameter
15 (13.8); snout length 8.8—9.5 (9.3); bony
interorbital width 8.4—8.8 (8.6); upper-jaw
length 18—20 (19.6); caudal-peduncle depth
14-16 (14.9); caudal-peduncle length 21-—
24 (19.4); first dorsal-spine length 3.0—3.6
(5.0); second dorsal-spine length 8.3—9.5
(9.3); third dorsal-spine length 19-20
(18.6); fourth dorsal-spine length 16—19
(18.6); spine in second dorsal fin 13-15
(13.8); first anal-spine length 3.1—4.1 (3.3);
second anal-spine length 12—15 (11.8); pec-
toral-fin length 21—24 (20.1); pelvic-fin
length 23-27 (23.4).
Dorsal fin VII-—I,9; anal fin II,8; pectoral
fin 14—14; pelvic fin 1,5; principal caudal
rays 9 + 8; simple pored lateral-line scales
24; transverse scale rows above lateral line
2; transverse scale rows below lateral line
5—6; median predorsal scales 2; circumpe-
duncular scale rows 12 (5+2+5). Total gill-
rakers 14-15 (14), well developed 9-10
(9), upper arch 2+1, lower arch 8—9+2-—3
(8+°3):
Villiform teeth in a wide band on the pre-
maxilla and dentary; two rows on the pal-
atine and vomer; none on ectopterygoid,
endopterygoid or basihyal.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Apogon gardineri, BPBM 24775, 42 mm SL, from Mauritius, off Flic en Flac by J. E. Randall.
Supramaxilla absent. Posttemporal
smooth on posterior margin. Preopercle ser-
rate on vertical and horizontal margins,
smooth on ridge. Infraorbitals smooth, pos-
terior ones with crenulations. Scales cte-
noid. Caudal fin weakly emarginate.
Life colors.—Based on a Ektachrome
transparency of the Mauritian specimen the
color pattern is as follows: Iris yellow; head
(without dark or pale spots, bars or stripes)
and body with a light tannish background;
a small brownish spot at the beginning of
pored lateral-line scales; a darker brownish
bar beginning at the base of first five dorsal
spines and extending to about level of pec-
toral fin, as a chevron, angled posteriorly;
a second fainter bar or blotch under last
four or five soft rays of second dorsal fin
reaching to or just below pored lateral line;
a full dark brownish mark (spot or bar-like)
on caudal peduncle; first dorsal fin with a
dark brownish bar extending over first five
spines and contiguous with bar on the body,
membrane behind the fifth to seventh spines
pale; second dorsal, anal and caudal fins
pale without spots, bars or stripes; pelvic fin
pale except for the last third of the first two
soft rays.
Preserved color pattern.—In 70% ethyl
alcohol: The holotype has no color remain-
ing. Regan (1908) reported: ‘‘Faint traces
of 3 dark vertical bars, the first below the
spinous dorsal, the second below the soft
dorsal, the third on the caudal peduncle.
Spinous dorsal blackish except posteriorly;
ventrals blackish at the tips’’. The Mauri-
tian specimens have a pale intestine, with a
few melanophores on the stomach, perito-
neum pale; first dorsal fin dark, with a mark
extending onto the body to about even with
the pectoral fin; a second saddle extends
downward from the posterior part of the
second dorsal fin base; a diffuse blotch on
the caudal peduncle; a faint cheek mark and
a spot behind the eye.
Distribution.—Known only from the
Mascarene Plateau.
Remarks.—Regan (1908) suggested that
A. gardineri was related to Apogon ban-
danensis Bleeker, 1854a. Apogon banda-
nensis has more gill-raker (total gillrakers
25-29) and has black stomach and intestine.
Smith (1961) synonymized A. gardineri
with Apogon nigripinnis Cuvier in Cuvier
& Valenciennes, 1828. The holotype of A.
nigripinnis is a different species with 16
pectoral-fin rays, 16 total rudiments and gill
rakers, 12 well-developed rakers (3+2 up-
per arch, 10+1 lower arch), two predorsal
scales, a deeper body (depth 42.9% stan-
dard length), soft dorsal and anal fins black-
ish and a uniform brownish body without
VOLUME 113, NUMBER 1
an obvious ocellated spot on the side of the
body (faded?) or basicaudal spot (faded?).
The holotype of Apogon thurstoni Day,
1888, shares all of the above-listed char-
acters of A. nigripinnis, 16 pectoral-fin rays,
16 total rudiments and gill rakers, two pre-
dorsal scales, a body depth of 43.4% of
standard length, but has 10 well developed
gill rakers (2+2 upper arch, 8+4 lower
arch) and an obvious ocellated spot on side
about midline. All of the Southern African
material specimens here as A. nigripinnis
have an ocellus on the side, but otherwise
have meristics and body depth similar to the
type: 15-16 pectoral-fin rays, 16—18 total
rudiments and gill rakers, 11—12 well-de-
veloped rakers (2—3+2 upper arch, 9-
10+ 1—4 lower arch), two predorsal scales,
a body depth of 42—48% of standard length.
Apogon taeniatus Cuvier in Cuvier &
Valenciennes, 1828, appears closely related
to A. nigripinnis and may also be confused
with A. gardineri. However, A. taeniatus
variably has an ocellated spot within the
first dorsal saddle above the pectoral fin, a
small basicaudal spot, faint stripes on body,
16—19 total rudiments plus gill rakers, 9—
12 well-developed rakers (2—4+1—2 upper
arch, 8—10+3-—5 lower arch), 14—15 pecto-
ral-fin rays, 3 predorsal scales and a greater
body depth of 40—45% standard length.
The holotype of Apogon suezii Sauvage,
1883, shares with A. taeniatus an ocellated
spot above the pectoral fin, 17 total rudi-
ments and gill rakers, 12 well-developed
rakers (2+2 upper arch, 10+3 lower arch),
15 pectoral-fin rays, three predorsal scales
and a body depth of 40.9% standard length,
but it has a small basicaudal spot within a
darkish bar.
The low gill-raker count, body shape and
color pattern of A. gardineri may cause
some confusion with Apogon timorensis
Bleeker, 1854b and two of its synomyms
Apogon fraxineus (Smith, 1961) and Apo-
gon striatus Fourmanior & Crosnier, 1964,
which differs in having a thin, dark cheek
line from the eye onto the preopercle, a
black stomach and intestine, 15—17, usually
259
16 pectoral rays and fewer developed (6—8)
and more rudiments (4-7) on the lower
arch. Fourmanoir and Crosnier’s name is an
unavailable name as a secondary homonym
of Apogon striatus (Radcliffe in Smith &
Radcliffe, 1912).
Apogon gardineri is found in deeper wa-
ter. It does not appear to have been reported
from any of the shallow-water collections
from Mauritius, Cargados Carajos or the
Seychelles in the past several decades.
Many of the species in this complex appear
to have continental distributions, for ex-
ample, Gon (1986, map 1). Apogon gardi-
neri represents an insular species endemic
to the Mascarene Plateau.
Apogon regani Whitley, 1951
Pies) 2 2ac55
Synonyms: Apogon punctatus Regan,
1908, preoccupied by Apogon punctatus
Klunzinger, 1880.
Material examined.—Syntypes: Apogon
punctatus BMNH — 1908.3.23.86-—89;
4(31.9—-65.4); Cargados Carajos; 39-55 m.
Other material: BPBM 35475; (45); Sey-
chelles, N. of Aride I., 4°10’S, 55°44’E;
R/V Tyro, Station 714, 2.4-m Agassiz trawl;
J. van der Land et al.; 19 Dec 1992; 55 m,
color photo; USNM 307706; (44); Indian
Ocean, Saya de Malha Bank, 10°16'00’S,
61°09’00"E; Vityaz Cr. 17; Sta 2810; 8 Jan
1989; 50-70 m; x-ray.
Comparative material.—See A. gardi-
neri.
Diagnosis.—A species of Apogon in the
subgenus Ostorhinchus with 15-17 pectoral
rays, 4—6 rows of small spots on the body,
pale stomach, intestine and peritoneum and
14—16 total gill rakers.
Description.—For general body shape
see Fig. 5. Range of proportions (as per-
centages of standard lengths, syntypes in
parentheses): greatest body depth 44 (40.7—
43.0); head length 37—42 (40.4—42.5); eye
diameter 10—13 (10.8—12.5); snout length
10-11 (9.4—10.6); bony interorbital width
9.6-9.8 (8.7—-9.7); upper-jaw length 23
260
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Bige 5:
(20.8—22.6); caudal-peduncle depth 17-19
(15.9-17.0); caudal-peduncle length 20—22
(18.5—20.8); first dorsal-spine length 3.7—
3.9 (3.0—4.1); second dorsal-spine length
10 (6.6—8.1); third dorsal-spine length 22—
23 (18.5—21.6); fourth dorsal-spine length
20—22 (17.9—20.3); spine in second dorsal
fin 16-18 (13.3-—14.5); first anal-spine
length 4.1—5.0 (3.0—4.1); second anal-spine
length 15-16 (12.0—13.3); pectoral-fin
length 21—23 (21.9); pelvic-fin length 29—
33° (25.7=27.7):
Dorsal fin VII—I,9; anal fin 1,8; pectoral
fin 15-15, 15-16, 16—16, or 17-17; pelvic
fin I,5; principal caudal rays 9 + 8; pored
lateral-line scales 24—25; transverse scale
rows above lateral line 2; transverse scale
rows below lateral line 6—7; median pre-
dorsal scales 2—3; circumpeduncular scale
rows 12(5+2+5). Total gillrakers 14-16,
well-developed 10—11, upper arch 2+2 and
lower arch 8—9+ 1-3.
Villiform teeth in a band on the premax-
illa, the outer row slightly larger; several
rows anteriorly, two rows on side of the
Apogon regani, BPBM 35475, 45 mm SL, from the Seychelles, N. of Aride I. by J. E. Randall.
dentary; one to two rows on the palatine,
two rows On vomer; none on ectopterygoid,
endopterygoid or basihyal.
Vertebrae 10 + 14. Five free hypurals,
one pair of slender uroneurals, three epur-
als, a free parhypural. Three supraneurals,
two supernumerary spines on first dorsal
pterygiophore. Basisphenoid present. Su-
pramaxilla absent. Posttemporal serrate on
posterior margin. Preopercle serrate on ver-
tical and horizontal margins, smooth on the
ridge. Infraorbitals smooth, the second
crenulate and third with a single spine.
Scales ctenoid. Caudal fin truncate.
Life colors.—Based on an Ektachrome
transparency of the Seychelles specimen the
color pattern is as follows: Iris pale; head
and body with a pale tannish background
on the upper parts; a dark brown oblique
line behind the eye; head silvery below eye
level; body silvery below the pored lateral
line on body to about the end of anal-fin
base, faint brownish spots on some, but not
all scales above the pored lateral line; larger
brownish spots on each scale below the
VOLUME 113, NUMBER 1
pored lateral line, forming about five lines
of spots, reducing to two lines on the an-
terior portion of the caudal peduncle; a
brownish area on the dorsum below the first
four or five spines; another mark on dorsum
below the fifth soft dorsal ray; caudal and
pectoral fins pale; first dorsal dusky and
whitish with a spot near the base of the
fourth to sixth spines; second dorsal fin
dusky; anal fin with a dark brownish edge
to the second anal spine and first soft ray,
the rest of fin dusky, pelvic fins dark
brownish except the fifth soft ray pale.
Preserved color pattern.—In 70% ethyl
alcohol: the syntypes with three to four
rows of small spots below the lateral line
on side of body. The pelvic fins and second
dorsal fin blackish. Stomach, intestine and
peritoneum pale. The other two specimens
with six rows of small spots on side of body
to caudal peduncle, one row above the lat-
eral line; three marks behind eye, a narrow
oblique cheek mark, a mark between the
eye and preopercle just below the mid-line
of the head, and an oblique mark past the
top of the preopercle; all vertical fins dark-
ish, caudal fin pale.
Distribution.—Known only from the
Mascarene Plateau.
Remarks.—Whitley (1951) created the
replacement name, A. regani for A. punc-
tatus Regan, 1908, which is preoccupied by
Apogon punctatus Klunzinger, 1880, a pri-
mary homonym. Klunzinger’s species is
presently placed in Vincentia, a mostly
warm temperate species group from Austra-
lia and Tasmania. According to the Inter-
national Commission on Zoological No-
menclature (1985: Art 59b) Regan’s hom-
onym is to be treated as permanently re-
jected.
Regan suggested a relationship with A.
nigripinnis. Smith (1961) synonymized A.
punctatus with A. nigripinnis. However, A.
regani differs from all known species in the
A. nigripinnis group by the presence of
small spots, one per scale, on the body. A.
regani represents an insular species in this
group.
261
This species is found in deeper water.
Randall & Egmond (1994) reported A. re-
gani as A. punctatus from the Seychelles
and provided a color photograph without
description. It does not appear to have been
reported from any of the shallow-water col-
lections from Mauritius or Cargados Cara-
jos in the past several decades. Apogon re-
gani represents the second insular species
endemic to the Mascarene Plateau in the A.
nigripinnis complex.
Acknowledgments
For the loan of material, the use of mu-
seum facilities and aid in curatorial pro-
cesses I thank: Arnold Y. Suzumoto
(BPBM), David Catania, Tomio Iwamoto,
William N. Eschmeyer (CAS), Susan L.
Jewett, David G. Smith, and Jeffery T. Wil-
liams (USNM). Jeffery T. Williams helped
with photography and David G. Smith pro-
vided x-ray photography of some speci-
mens. Many other colleagues over the past
two decades at various museums have pro-
vided valuable assistance in my studies of
the Apogonidae. Leonard P. Schultz funds
were provided by Victor G. Springer
(USNM) for several study trips to the
Smithsonian. John E. Randall provided col-
or transparencies of A. gardineri and A. re-
gani. Ofer Gon and John E. Randall re-
viewed an earlier draft of this manuscript.
Two anonymous reviewers provided helpful
comments.
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):264—268. 2000.
A new species of Pogonophryne
(Pisces: Perciformes: Artedidraconidae) from East Antarctica
Richard R. Eakin and Arcady V. Balushkin
(RRE) Department of Life Sciences, University of New England, Westbrook College Campus,
Portland, Maine 04103; (AVB) Zoological Institute, Russian Academy of Sciences,
Universitetskaya nab. 1, St. Petersburg, 199034 Russia
Abstract.—A new species of notothenioid fish, Pogonophryne squamibar-
bata, is described from East Antarctica. This dorsally-spotted species belongs
to the ‘“‘mentella’’ group of the genus and is characterized by having a relatively
short mental barbel (16% SL) with bluntly palmate, overlapping, scale-like
processes on its terminal expansion, a wide interorbital region (6.9% SL), and
a long opercular-subopercular distance (19.2% SL).
German research cruises of the R/V Po-
larstern in Antarctic waters have yielded
several new species of the artedidraconid
genus Pogonophryne (Eakin 1987, Balush-
kin & Eakin 1998, Eakin & Balushkin
1998). The “‘mentella’’ group is the most
speciose of the genus, comprising at least
10 species, three of which have been re-
cently described (Balushkin & Eakin 1998,
Eakin & Balushkin 1998, Eakin & Eastman
1998). This paper continues our descrip-
tions of new species of Pogonophryne in
the Zoological Museum collections at Ham-
burg University.
Measurements and counts follow those of
Balushkin & Eakin (1998) and Eakin &
Eastman (1998). Opercular-subopercular
distance is measured from the posterior
edge (hook) of the opercular bone to the
anterior edge of the subopercular bone.
Measurements are presented as percentages
of standard length unless otherwise indicat-
ed. ISH refers to the Zoological Museum
collections (formerly Institut fiir Seefischer-
ei) currently maintained at Hamburg Uni-
versity (ZMH).
Pogonophryne squamibarbata,
new species
Figs. 1-2
Holotype.—ISH 65-1991; male, 147 mm
SL; R/V Polarstern station 211, East Ant-
arctica (69°58.9'S, 05°8.4’E), 651-742 m;
10 Mar 1991.
Diagnosis.—This species of Pogono-
phryne belongs to the “‘mentella”’ group.
The relatively short (16% SL) mental bar-
bel has a terminal expansion composed of
bluntly palmate, flattened processes ar-
ranged in an overlapping, scale-like pattern.
This species also has a wide interorbital re-
gion (6.9% SL) and a long opercular-sub-
opercular distance (19.2% SL).
Description.—Body robust anteriorly, ta-
pering to low, narrow caudal peduncle.
Head length 43.5; posttemporal ridges not
well developed (upper limb of posttemporal
bone not arched upward to form a promi-
nent curved ridge), depth of head at this
level 19.7; head width at preoperculars
37.4. Body depth at second dorsal fin origin
17.3; body depth at anal fin origin 13.6;
body width at anal fin origin 10.5; caudal
peduncle depth 6.5. Snout broadly rounded
in dorsal view; internostril distance 9.2;
snout length 8.6; eye filling orbit; diameter
of orbit 9.9. Interorbital region wide, bony
measurement 6.9. Opercular-subopercular
distance long, 19.2. Postorbital length of
head 23.7. Jaw width at posterior ends of
maxillaries 25.5. Upper jaw length 19.3;
posterior end of maxillary extending slight-
ly beyond pupil of eye; lower jaw project-
VOLUME 113, NUMBER 1
male, 147 mm SL.
>
Pogonophryne squamibarbata, holotype, ISH 65-1991
Fig. 1:
N
Nn
266
ing beyond upper a distance 7.8 of head
length. Tongue long, extending anteriorly
slightly in front of tip of upper jaw. Upper
jaw with two rows of teeth (largest near
symphysis) in a broad arc. Lower jaw teeth
in two rows near symphysis, becoming uni-
serial posteriorly (nine large teeth on both
sides) in a V-shaped pattern. Mental barbel
(Fig. 2) rather short (16% SL) and with a
terminal expansion composed of bluntly
palmate, flattened processes arranged in an
overlapping, scale-like pattern and continu-
ing onto stalk; barbel length 16.0; terminal
expansion 34.0 of barbel length and slightly
wider than stalk. Anterior gillrakers on first
arch 2 + 0 + 7 = 9; posterior gillrakers O
+ 1+ 7 =s3tonla7
First antedorsal distance 37.4; second an-
tedorsal distance 46.3; anteanal distance
64.6. Length of second dorsal fin base 52.4;
length of anal fin base 33.0; length of cau-
dal fin 25.3;. length of pectoral fin 28:7:
width of pectoral fin base 10.5; length of
pelvic fin 20.4. Interdorsal distance 7.5.
First dorsal fin with two spines of about
equal length, 9.2. Second dorsal fin with 27
rays; length of longest (fifth) ray 15.3. Ratio
of longest first dorsal fin spine to longest
second dorsal fin ray 0.60. Anal rays 17.
Pectoral rays 20. Upper lateral line with
about 26 (left) and 27 (right) pores (tubular
scales), ending under the twenty-first ray of
the second dorsal fin. Middle lateral line
with about 11 tubular scales (13 pores) on
the left side and 11 tubular scales on the
right side; tubular scales originate under the
eighteenth ray of the second dorsal fin. Ce-
phalic lateral-line pores typical for Pogon-
ophryne: preoperculo-mandibular canal
with nine pores; infraorbital canal with sev-
en pores; supraorbital canals with two nasal
pores, two interorbital pores, and one un-
paired coronal pore; temporal canal with six
pores; and supratemporal canal incomplete
across occiput (one pore on each side). Ver-
tebrae 16 + 21 = 37 (second preural ver-
tebra appears double, being longer than the
others and possessing two neural and two
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
hemal spines, but is counted as one ele-
ment).
Radiographic analysis of skeleton.—
Number of vertebrae to first pterygiophore
of first dorsal fin 2; to first pterygiophore
of second dorsal fin 6; to first pterygiophore
of anal fin 16. Free caudal vertebrae count-
ing from last pterygiophore of second dor-
sal fin 6; counting from last pterygiophore
of anal fin 6. Four pterygiophores anterior
to first pterygiophore of second dorsal fin;
the anterior two support spines of the first
dorsal fin, while the posterior two are ray-
less pterygiophores. One pterygiophore an-
terior to first caudal vertebra under abdom-
inal section of vertebral column. Four hy-
pural plates attached to urostyle; parhypural
(PH) + lower complex hypural (H1 + H2)
+ upper complex hypural (H3 + H4) +
minimal hypural (H5). All hypural plates
except H5 fused to urostyle. Caudal fin with
21 rays (4 upper procurrent + 13 principal
+ 4 lower procurrent). Distribution of prin-
cipal caudal rays with reference to hypural
plates: 1-6—4—2. Epurals 3, the anterior two
of which resemble in shape the neural spine
of the first preural vertebra. One pair of uro-
neurals.
Color in alcohol.—Head and body with
brownish markings dorsally and laterally on
yellowish background. Markings on head in
the form of round spots and vermiculations;
those on body larger and more irregular. A
broad, dark horizontal marking extends
along base of anal fin. Ventral surfaces of
head and body dusky and unmarked. First
dorsal fin light; second dorsal fin with dark
oblique stripes. Anal fin light. Caudal fin
with about six dark vertical stripes. Pectoral
fins with about six dark vertical stripes. Pel-
vic fins light. Mental barbel light.
Etymology.—Named for the overlapping,
scale-like processes on the mental barbel
(Latin sqguama: a scale; barb: a beard).
Comparative notes.—The new species
belongs to the “‘mentella’’ group of the ge-
nus which is generally characterized by
having a well developed mental barbel
(elongation, complex terminal expansion,
VOLUME 113, NUMBER 1 267
Fig. 2. Mental barbel of Pogonophryne squamibarbata, holotype; a, terminal expansion (10); b, detail of
terminal expansion showing overlapping, palmate processes (X40).
268
or both), relatively undeveloped posttem-
poral ridges (upper limb of posttemporal
bone not arched upward to form a promi-
nent curved ridge), and large, sparsely dis-
tributed, round spots and vermiculations on
the dorsal and lateral surfaces of the head
(Balushkin & Eakin 1998). The mental bar-
bel somewhat resembles that of P. mentella
in having a terminal expansion composed
of bluntly palmate, flattened processes.
However, the barbel of P. squamibarbata is
much shorter (16% SL) than that of the ho-
lotype of P. mentella (27.1% SL; Andria-
shev 1967) and has a relatively longer ter-
minal expansion (34% of barbel length
compared to 26.4% of barbel length; mea-
sured from Andriashev 1967, Fig. 1). The
scale-like pattern of densely packed, over-
lapping processes on the mental barbel of
P. squamibarbata is unlike that of any other
species of Pogonophryne. This species also
has a wide interorbital region (6.9% SL)
and a long opercular-subopercular distance
(19 29% SE):
Acknowledgments
We thank Horst Wilkens of the Zoolog-
ical Museum, Hamburg University, for his
kind invitation to examine the ISH collec-
tions and for his hospitality during our visit
in June 1997. RRE thanks the University of
New England/Westbrook College for partial
financial support from a Bingham Faculty
Enrichment Grant. AVB is grateful for fi-
nancial support from the German Scientific
Society (Grant no. 436 RUS-17/41/97). The
work was also supported by the Russian
Foundation for Basic Research (Grant no.
97-04-48950). We are indebted to Gudrun
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Schulze of the Zoological Museum, Ham-
burg University, for her help in all aspects
of our work. We thank Jacob Levenson of
the University of New England for photo-
graphing the holotype and Helga Kapp of
the Zoological Museum, Hamburg Univer-
sity, for help and use of equipment in pho-
tographing the mental barbel. We thank An-
gelika Brandt of the Zoological Museum,
Hamburg University, for checking collec-
tion data. We also thank Joseph Eastman of
Ohio University for critically reading the
manuscript.
Literature Cited
Andriashev, A. P. 1967. A review of the plunder fishes
of the genus Pogonophryne Regan (Harpagifer-
idae) with descriptions of five new species from
the East Antarctic and South Orkney Islands.
Pp. 389-412 in A. P. Andriashev and P. V.
Ushakov, eds., Biological results of the Soviet
Antarctic Expedition (1955-1958), vol. 3. Len-
ingrad Zoological Institute, Academy of Scienc-
es of the USSR.
Balushkin, A. V., & R. Eakin. 1998. A new toad plun-
derfish Pogonophryne fusca sp. nova (Fam. Ar-
tedidraconidae: Notothenioidei) with notes on
species composition and species groups in the
genus Pogonophryne Regan.—Journal of Ich-
thyology 38:574—579 (English translation from
Voprosy Ikhtiologii 38:598—603).
Eakin, R. R. 1987. Two new species of Pogonophryne
(Pisces, Harpagiferidae) from the Weddell Sea,
Antarctica.—Archiv fiir Fischereiwissenschaft
38:57-74.
, & A. V. Balushkin. 1998. A new species of
toadlike plunderfish Pogonophryne orangiensis
sp. nova (Artedidraconidae, Notothenioidei)
from the Weddell Sea, Antarctica.—Journal of
Ichthyology 38:800—803 (English translation
from Voprosy Ikhtiologii 38:830—833).
, & J. T. Eastman. 1998. New species of Po-
gonophryne (Pisces, Artedidraconidae) from the
Ross Sea, Antarctica.—Copeia 1998:1005-—
1009.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):269—283. 2000.
A new genus and species of inseminating fish
(Teleostei: Characidae: Cheirodontinae: Compsurini) from
South America with uniquely derived caudal-fin dermal papillae
Luiz R. Malabarba and Stanley H. Weitzman
(LRM) Departamento de Zoologia - IB, Universidade Federal do Rio Grande do Sul,
Av. Paulo Gama s/n, 90046-900, Porto Alegre, RS, Brasil, and Museu de Ciéncias e Tecnologia,
PUCRS, Av. Ipiranga 6681, 90619-900, Porto Alegre, RS, Brazil; (SHW) Division of Fishes,
Department of Vertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560-0159,
U.S.A.
Abstract.—Kolpotocheirodon theloura, a new genus and species of the chei-
rodontine tribe Compsurini, is described from the uppermost tributaries of the
rio Sao Francisco and rio Parana in central Brazil. Males of the new species
bear a caudal-fin organ formed by fin-ray hooks and hypertrophied elongate
dermal flaps along the fin rays, mostly in the ventral lobe of the caudal fin.
The largest flap, attached dorsally to the nineteenth principal caudal-fin ray
(ventral principal fin ray of the ventral caudal-fin lobe), is bordered with a
series of tabs, and extends dorsally over most of the flaps extending from
caudal-fin rays 17 and 18. There are also a series of fleshy papillae distributed
along the border of the remaining ventral caudal-fin ray flaps or along the fin
rays just dorsal to the flap-bearing fin rays. These tabs and papillae are un-
known in both other inseminating and all externally fertilizing characids and
therefore are considered derived. The new species is referred to the Compsurini
of the Cheirodontinae on the basis of a cladistic diagnosis of these taxa. The
relationships of K. theloura with other inseminating compsurins bearing mod-
ified caudal fins are discussed.
Resumo.—Kolpotocheirodon theloura, género e espécie novos da tribo
Compsurini de Cheirodontinae, é descrito para os tributarios superiores do rio
Sao Francisco e rio Parana no Planalto Central do Brasil. Machos da nova
espécie possuem um 6rgao formado por ganchos nos raios da nadadeira caudal
e por dobras de pele (‘“‘flaps’’) hipertrofiadas ao longo dos raios, principalmente
no lobo ventral da nadadeira caudal. O maior “‘flap’’, conectado ao longo da
face dorsal do 19° raio da nadadeira caudal (raio principal inferior do lobo
ventral da nadadeira caudal), é marginado por uma série de tabiques, e se
estende dorsalmente sobre as dobras de pele originadas nos 17° e 18° raios.
Existe ainda uma série de papilas carnosas distribuidas ao longo das demais
dobras de pele do lobo ventral da nadadeira caudal ou ao longo dos raios
imediatamente superiores aos raios com dobras de pele. Estes tabiques e papilas
sao desconhecidos em outros caracideos, tanto naqueles com inseminagao como
nos com fertilizagao externa, sendo considerados como caracteres derivados.
A espécie nova é referida aos Compsurini em Cheirodontinae com base em
uma diagnose cladistica destes taxons. SAo discutidas as relagdes de K. theloura
com outros compsurineos de nadadeiras caudais modificadas.
The new genus and species herein de- with modified, possibly glandular tissues,
scribed (see Figs. 1-3) is a cheirodontine and hooks on the caudal fin of males. It was
270
discovered in the collections of the Museu
de Zoologia, Universidade de Sao Paulo by
one of us (L.R.M.) while reviewing chei-
rodontine characids of southeastern and
eastern Brazil. The species was first hy-
pothesized to belong to a cheirodontine
clade diagnosed by specialized modified
anal-fin hooks and derived scales, fin rays
and/or hypertrophied soft tissues on the
caudal fin. In the unpublished thesis of Mal-
abarba (1994), this clade includes Sacco-
derma Schultz, Compsura Eigenmann, Ma-
cropsobrycon Eigenmann, Acinocheirodon
Malabarba & Weitzman, and the Central
American ‘“‘Odontostilbe’’ dialeptura (Fink
& Weitzman) and “‘O.”’ mitoptera (Fink &
Weitzman). Later, the new taxon (listed as
undescribed genus and species A in Burns
et al. 1997) and all cheirodontine taxa listed
above (see Burns et al. 1997) were found
to be inseminating species. Subsequently,
this putative clade was recognized as a
cheirodontine tribe, the Compsurini (Mala-
barba, Weitzman, & Burns in Malabarba
1998), including the taxon described herein
(therein referred as new genus and species
A). The significance of these characters in
resolving the phylogeny of compsurin chei-
rodontines, as well as its possible homology
to similar characters found in glandulocau-
dine characids, were further discussed by
Malabarba & Weitzman (1999).
Males of the new genus and species have
hooks on the caudal fin (Fig. 4) and hyper-
trophied dermal flaps along the caudal-fin
rays (Fig. 5). The largest flap attached dor-
sally to the 19th ventral lobe caudal-fin ray
is bordered with a series of tabs, and ex-
tends dorsally over most of the flaps ex-
tending from caudal-fin rays 17 and 18.
There is also a series of fleshy papillae dis-
tributed along the border of the remaining
ventral caudal-fin ray flaps or along the fin
rays just dorsal to the fin rays bearing flaps.
These tabs and papillae are unknown in the
remaining inseminating cheirodontines, as
well as on other inseminating or externally
fertilizing species of characids and are con-
sequently hypothesized to be derived. This
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
is the only species of the Compsurini
known to have aquasperm (a nearly spher-
ical or spherical sperm nucleus, similar to
that found in externally fertilizing chara-
cids; see Burns et al. 1997:434, fig. 1A).
All other species of the Compsurini so far
investigated have elongate sperm cell bod-
ies (see Burns et al. 1997:434, fig. 1B—H).
Methods
The systematic methods, counts and
measurements used here are the same of
those described and used by Malabarba &
Weitzman (1999). The following acronyms
are used for institutions and collections:
MCP, Museu de Ciéncias e Tecnologia,
Pontificia Universidade Catdlica do Rio
Grande do Sul, Porto Alegre; and MZUSP,
Museu de Zoologia da Universidade de Sao
Paulo, Sao Paulo. The following abbrevia-
tions are used in the text or figures: SL =
standard length; HL = head length; c&s =
alizarin red s and alcian blue stained spec-
imens cleared with trypsin; spm(s) = spec-
imen(s). The comparative material is the
same listed in Malabarba & Weitzman
(1999).
Kolpotocheirodon, new genus
Type species.—Kolpotocheirodon thel-
oura, new species.
Diagnosis.—The following apomorphies
diagnose Kolpotocheirodon:
A specialized caudal organ (Fig. 5) is
present at the proximal region of the ventral
caudal-fin lobe of the males (character
number 36 in Malabarba 1998). This con-
sists of hypertrophied elongate dermal flaps
attached along the fin rays and a series of
relatively flat tabs attached along the ex-
posed border of the largest flap. The base
of this flap is attached along the anterior
approximately one-third of the nineteenth
caudal-fin ray. Visually moving dorsally,
the flaps decrease in length and width until
those of the thirteenth or fourteenth fin rays
are relatively short, narrow, and almost un-
detectable. Each flap, other than that of the
VOLUME 113, NUMBER 1
nineteenth ray, bears papillae in a single se-
ries along its exposed border. The tabs of
the nineteenth fin ray flap extend dorsally
while the papillae of the fourteenth to at
least the sixteenth or seventeenth flap are
directed ventrally.
Hooks on the anal-fin rays of mature
males (Fig. 6) occur along the posteriola-
teral border of the posterior unbranched and
five anterior branched anal-fin rays (Char-
acter 30 in Malabarba 1998). Although the
number of hook-bearing anal-fin rays in any
species of the Characidae with hooks is var-
iable according to the degree of maturation,
it seems to be constant in fully mature spec-
imens of some species. All fully mature
males of Kolpotocheirodon theloura have
fully developed hooks restricted to six of
the anterior anal-fin rays. The remaining
species of the Compsurini have hooks on a
larger number of anal-fin rays, with the ex-
ception of the species of Saccoderma,
which have anal-fin hooks only along the
posterior ray of the anterior unbranched
rays and the four most anterior branched
anal-fin rays.
The twelfth and thirteenth caudal-fin rays
are somewhat curved, being noticeably
concave along their dorsal borders at about
their basal half lengths (Fig. 4) and some-
what convex along their ventral borders
where the segments are slightly expanded
longitudinally (character 34, state 2 in Mal-
abarba 1998). This feature separates K.
theloura from other cheirodontines. Acin-
ocheirodon melanogramma Malabarba &
Weitzman (1999), another compsurin, also
has the basal halves of the caudal-fin rays
dorsally concave with ventrally expanded
segments, but in this species they are thir-
teenth and fourteenth, rather than twelfth
and thirteenth rays.
Etymology.—The first component of the
name Kolpotocheirodon is from the Greek,
kolpotos = formed into folds, and refers to
the caudal organ formed by hypertrophied
dermal folds along the caudal-fin rays. The
second component refers to the characid ge-
nus Cheirodon Girard, in reference to our
271
referral of this new genus to the Cheiro-
dontinae.
Kolpotocheirodon theloura, new species
Figs. 1-3
All specimens from Brazil, Brasilia, Dis-
trito Federal, except when noted.
Holotype.-—MZUSP 55194, 1 male, 27.3
mm SL, small marsh at Curva da Morte,
Goias, 3 May 1978, E. K. Bastos.
Paratypes.—Rio Parana tributaries: MCP
1161, Lt male 25-9 mm SL; eds; MZUSP
38840, 2 females, 27.9 and 29.7 mm SL,
and MZUSP 38839, 1 male, 29.9 mm SL,
ribeirao do Gama, just above the mouth of
ribeirao Taquara, 1 Oct 1985, J. P Viana
Mendes. MZUSP 39014, 1 female, 24.0
mm SL and MCP 11160, 1 female, 22.2
mm SL, ribeirao Riacho Fundo, tributary of
rio Sao Bartolomeu, 3 Jul 1985, M. Ribeiro.
MZUSP 42802, 1 male, 27.8 mm SL, 1 fe-
male, 24.6 mm SL, collected with the ho-
lotype. Rio Sao Francisco tributaries: MCP
12204, 4 spms. (1 alcohol, 3 c&s), 14.0—
16:0:mm SL, and MZUSP 35722; 12 spms,
14.7—19.5 mm SL, corrego Vargem de Tras,
1—2 Apr 1979, N. A. Menezes & E. K. Bas-
tos. MZUSP 42801, 2 males, 22.9—23.8
mm SL and 1 female, 20.2 mm SL, lagoa
Feia, 3 May 1978, E. K. Bastos.
Diagnosis.—The same as for the genus.
Distinguishing characters.—The pres-
ence of spherical sperm nuclei (aquasperm;
see Burns et al. 1997:434, fig. 1A and tab.
1, “undescribed genus and species A’’) is
plesiomorphic for the Compsurini, accord-
ing to the parsimony analysis in Malabarba
(1998). Its presence in K. theloura distin-
guishes this species from all other species
of the Compsurini so far investigated,
which have elongated sperm nuclei (see
Burns et al. 1997:434, fig. 1B—H and tab.
1).
Kolpotocheirodon theloura also has an
atypical color pattern for the Cheirodonti-
nae of three to five very small vertical bars
on the sides of the body, crossing the nar-
row lateral horizontal body stripe. These are
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig, <1:
at Curva da Morte, Goias, Brazil.
located dorsal to the visceral cavity, and be-
tween the pseudotympanum and the area
ventral to the dorsal fin (Figs. 1-3).
Kolpotocheirodon theloura can also be
distinguished from other cheirodontines by
a conspicuous dark brown band formed of
a series of spots distributed along all of the
hook-bearing portion of the anal-fin of the
males. These pigment spots are placed at
approximately the midlength of the poste-
riormost of the anterior unbranched anal-fin
rays and the five anteriormost branched
anal-fin rays. The most conspicuous and
darkest spot is at the anterior surface of the
first branched ray (Fig. 1; although also pre-
sent in the paratypes, the dark brown spot
cannot be seen in the black ground Figs. 2—
3). A similar, but less conspicuous band of
spots occurs in females along the midlength
of the anal-fin rays. These are more strong-
ly pigmented in the anterior portion of the
fin.
Description —Morphometric data sum-
marized in Table 1.
Body moderately elongate and com-
pressed, greatest depth at dorsal-fin origin.
Predorsal profile convex, slightly concave
in region of supraoccipital process. Profile
of body from base of posterior dorsal-fin
ray to origin of adipose fin straight or mod-
erately convex. Ventral profile of body con-
vex from tip of lower jaw to pelvic-fin or-
igin and moderately concave from there to
anal-fin origin. Body profile along anal-fin
Kolpotocheirodon theloura, new species, holotype, male, MZUSP 55194, SL 27.3 mm; small marsh
base of males moderately concave in ante-
rior half and convex posteriorly. In female
entire fin base relatively straight. Dorsal
and ventral profile of caudal peduncle con-
cave. Largest mature male examined with
an expanded and rounded dorsal and ventral
caudal-peduncle profiles. Caudal peduncle
about as long as deep or somewhat shorter
than deep.
Head small and snout short, rounded.
Mouth terminal. Maxilla short, positioned
at an angle of approximately 45 degrees rel-
ative to long axis of body. Posterior extrem-
ity of maxilla reaching vertical that passes
through anterior border of eye.
Premaxilla with 4 (rarely 5) symmetrical
teeth having 9-11 small evenly spaced
cusps all about equal in size (Fig. 7). Cut-
ting edge arched in most medial tooth and
almost straight in most lateral tooth. Max-
illa with 2 (rarely 3) teeth, similar in form
to those of premaxilla, with 7—11 cusps.
Cutting edge slightly arched to almost
straight. Dentary with 4 large teeth with 5,
7, or 9 cusps; followed by 1 or 2 smaller
teeth with 3, 5 or 7 cusps. Teeth following
second tooth asymmetrical with most lateral
cusp situated towards tooth base and most
medial cusp more distally located. Cusps
small and regular and approximately equal
in size. Cutting edge slightly arched to al-
most straight.
Dorsal-fin rays, ii, 9, n = 29. First un-
branched ray about half length of second.
VOLUME 113, NUMBER 1
Dorsal-fin origin approximately at mid-
length of body. Proximal portion of dermal
flaps along largest unbranched and first
branched dorsal-fin rays bordered by series
of papillae in both sexes. Adipose-fin origin
at, or slightly anterior to, vertical through
insertion of posteriormost anal-fin ray.
Anal-fin rays, iii, 18, Gi-iv, X = 3.3, 17—
fee = 19.1, mn = 22). Anal-fin origin
slightly posterior to vertical passing through
base of posteriormost dorsal-fin ray. Anal
fin of females with anterior 5—6 branched
rays very long, forming prominent anterior
lobe. Rays posterior to anterior lobe abrupt-
ly shorter and distal border of anal fin con-
cave posterior to anterior lobe. Distal bor-
der of male anal fin decreasing in length
gradually so that although anterior lobe
larger than but not as abruptly distinct as in
females. Anal-fin rays of males with slen-
der, elongate retrorse hooks on longest un-
branched ray, and anterior first 5 branched
rays, usually with 4 to 5 segments bearing
hooks on each ray (Fig. 6). Hooks inserted
at posterolateral border of fin rays, bent
over lateral surface of fin ray and anteriorly
directed. Hooks located on _ posterior
branches of rays only; never on proximal
portions of main body of rays. One, rarely
two, bilateral pair of bony hooks per ray
segment.
Pectoral-fin rays, i, 9, (i, 8-11, X = 9.2,
n = 28). Distal ends of longest rays extend
slightly beyond pelvic-fin origin. Pelvic-fin
fet i 78 = F1, ) = (29). Pelvic-
fin origin anterior to vertical passing
through dorsal-fin origin. Tip of fin reach-
ing anal-fin origin in males but not in fe-
males. Male pelvic fins with unilateral, ven-
tromedial retrorse bony hooks (Fig. 8);
branched rays bearing one, rarely two slen-
der hooks per segment along most of
branched ray lengths, including both pri-
mary branches and often posterior second-
ary branches. Distal sections of unbranched
part of each branched ray with one to oc-
casionally few hooks. Number of hooks
varies among different males, those males
bearing more hooks appearing more mature
Kolpotocheirodon theloura, new species, paratype, male, MZUSP 42802, SL 27.8 mm; small marsh at Curva da Morte, Goids, Brazil.
Fig. 2.
274
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3.
at Curva da Morte, Goias, Brazil.
when considering other secondary sexual
characters such as caudal-fin papillae.
Principal caudal-fin rays 10/9, n = 29.
Proximal portion of thirteenth or fourteenth
through nineteenth caudal-fin rays in ven-
tral caudal fin lobe with hypertrophied der-
mal flaps. Largest flap extending dorsally
from nineteenth principal caudal-fin ray
(Fig. 5). Each flap bordered by one series
of external papillae, more numerous and
more developed in males. Proximal portion
Kolpotocheirodon theloura, new species, paratype, female, MZUSP 42802, SL 24.6 mm; small marsh
of third caudal-fin ray in dorsal caudal-fin
lobe with hypertrophied dermal flaps and
external papillae only in males. Males with
modified 12th to 14th or 15th principal cau-
dal-fin rays; these rays bearing (on each
side) row of 4—6 dorsoanteriorly directed
hooks along dorsal divisions (Fig. 4). Dor-
sal and ventral procurrent rays equivalent in
numbers and structure in both sexes, similar
to those of most characids. Dorsal procur-
rent caudal-fin rays, 13 (9-13, X = 10.8, n
Table 1.—Morphometrics of Kolpotocheirodon theloura, new species. Standard length is expressed in mm;
measurements through head length are percentages of standard length; the last four entries are percentages of
head length. Range includes the holotype, MZUSP 55194, and following paratypes MZUSP 38839, MZUSP
42801, MZUSP 42802, MCP 11161, MZUSP 38840, MZUSP 39014, MCP 11160.
Males
Holotype n Low
Standard length (mm) DS 6 2pe9
Snout to anal-fin origin 62.3 6 60.1
Snout to dorsal-fin origin 49.5 6 48.9
Snout to pelvic-fin origin 45.4 5 41.7
Dorsal-fin base length hg Ws 5) 7
Anal-fin base length 29.3 5 26.4
Caudal peduncle length of 5 2h)
Caudal peduncle depth 14.7 5) 29
Depth at dorsal-fin origin 5525 5 32.4
Dorsal-fin height 28.2 =) 27.0
Pelvic-fin length 20:9 5 19.4
Pectoral-fin length 23.8 5 2055
Bony head length 26.0 5 24.4
Snout length 13:3 3) 18.3
Upper jaw length 25.4 4 25.4
Horizontal eye diameter Sez 5 32.4
Least interorbital width 29.6 5 29.6
Females
High XK nen \Ebote Pauklich ee ose
259 26.3 5) 2ae2 294] Dae
63.9 61.8 3) 62:2 66.3 65.2
S15 50.0 5 48.3 5310 5 kat
46.5 44.9 5 44.1 45.5 44.6
13-1 [25 5 12.6 14.0 13,3
29.4 28.6 5 26.2 28.5 27.4
12.4 11.0 5 11.0 12-2 11.6
14.7 Le 5 LAS 13:3 122
56:1 34.3 5 30.4 37.6 34.4
30.4 233, 5 Doe 29.3 27.9
2, Wee 20.4 5 15.0 18.6 ig ee |
23.8 PMs?) + 19.5 23:3 21.4
26.8 25-8 5 Play, yada 26.5
24.1 222 5 2A 24.7 22.9
29.2 213 2 297 32:2 31.0
39.1 36.1 5 35.6 38.5 36.8
36.2 31.3 5) 29.2 IEA 30.6
VOLUME 113, NUMBER 1
Fig. 4. Kolpotocheirodon theloura, new species, paratype, MCP 11161, c&s male, SL 25.9 mm. Caudal-fin
rays eleven (11th) through fifteen (15th) of the ventral caudal-fin lobe, other lower caudal-fin rays only partially
indicated. Figure illustrates the position and number of caudal-fin ray hooks along fin rays twelve to fourteen.
Anterior is to left.
= 5). Ventral procurrent caudal-fin rays, 9
(n = 5).
Scales cycloid, moderately large. Lateral
line poring incomplete, 8, (6-10, X = 7.6,
n = 21) anterior perforated scales. Scales
in lateral series 33, (31-34, X = 32.8, n =
19). Scale rows between dorsal-fin origin
mueeeneral line 5, (5-6, X — 5.1, n — 22).
Scale rows between lateral line and pelvic-
famien 4, G—4, X = 3:6; n = 22). Pre-
dorsal scales, when in regular series 11
esti oX = 10.8,.n = 9). Scale. rows
around caudal peduncle 14 (nm = 5). Scale
Sheath on anal-fin base consisting of 3—5
scales covering unbranched rays and ante-
rior 4—5 branched rays.
Saoraneurals, 5° (3—5,°X' ="4.3).n' = 8).
Precaudal vertebrae, 15 (15-16, X = 15.1,
n = 12). Caudal vertebrae, 17 (17-18, X =
17.5, n = 12).
Color in alcohol.—Head and body pale
brownish yellow. Black lateral body stripe
very narrow, pale on caudal peduncle and
anterior to dorsal-fin origin (Figs. 1—3). Lat-
eral body stripe crossed by 3—5 very small
vertical bars between pseudotympanum and
area ventral to dorsal fin. Humeral spot ab-
sent. Caudal-fin base and posterior termi-
nation of caudal peduncle bear dark black,
vertically-elongate, lozenge-shaped spot
that reaches dorsal and ventral margins of
caudal peduncle. Caudal spot extends just
to most proximal portion of middle caudal-
fin rays otherwise white or pale brownish
yellow. All other fins hyaline except anal
fin. Anal fin of males with conspicuous
dark brown band distributed along all hook-
bearing portions of fin (approximately mid-
dle length of last unbranched anal-fin ray
and five anteriormost branched anal-fin
rays). Pattern most conspicuous at anterior
surface of first branched ray. Less conspic-
uous similar band of spots occurs in fe-
males, along midlength of most anterior
branched anal-fin rays.
Sexual dimorphism.—The females lack
hooks on the pelvic, anal, and caudal fins,
while the males bear numerous small hooks
on the pelvic and anal-fin rays. The male
holotype and paratypes bear about 4—6
hooks on caudal-fin rays 12—14 with more
of these hooks on ray 12 than on ray 14.
Both sexes bear dermal tabs and papillae on
the caudal fin, but the females lack these
papillae in the dorsal caudal-fin lobe al-
though they are present in males. The fe-
males have the distal border of the anal fin
with a distinct anterior lobe that is abruptly
curtailed at about branched rays 6—7. The
male distal anal-fin profile is not lobate. In-
stead the fin rays gradually diminish in
length from the anterior to posterior termi-
nations of the fin. The pelvic fins are longer
in the males (19.4—21.7% of SL) than in
the females (15.0—18.6% of SL), reaching
past the anal-fin origin in the males, but not
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 5.
portion of the ventral caudal-fin lobe. Figure illustrates the derived hypertrophied flaps bearing tabs and papillae
along the caudal-fin rays. White arrow indicates largest flap (1f) extending dorsally from the nineteenth principal
caudal-fin ray (19th), bordered by one series of flat tabs. All other flaps are bordered by one series of fleshy
papillae.
females. The relative position of the anal fin
also seems to be sexually dimorphic, with
the snout to anal-fin origin distance in
males (60.1—63.9% of SL) shorter than in
females (62.2—66.3% of SL). However, our
sample size is small and the examination of
larger population samples is necessary to
confirm this pattern of difference between
males and females.
Distribution.—Kolpotocheirodon thel-
oura has an interesting distribution, along
the uppermost tributaries of both the rio Sao
Francisco and the upper rio Parana basins,
near Brasilia, in central Brazil. It is found
close to an area called Aguas Emendadas
(joint waters), in allusion to the close prox-
imity of the head waters of the upper trib-
utaries of the rio Sao Francisco, alto rio Pa-
rana and rio Tocantins. This may explain
Kolpotocheirodon theloura, new species, holotype, male, MZUSP 55194, SL 27.3 mm. Proximal
the distribution of this species in two of
these major basins. So far, this species is
unknown from the rio Tocantins tributaries,
but samples from that area are too rare to
assure its absence in that drainage.
Etymology.—The name theloura is from
the Greek thele meaning nipple and oura
meaning tail. The name refers to the pres-
ence of papillae on the ventral lobe of the
caudal fin. The name is a noun in apposi-
tion.
Discussion.—The relationships of Kol-
potocheirodon theloura to other cheirodon-
tine characids is supported by the presence
of all four synapomorphies diagnosing the
Cheirodontinae (Malabarba, 1998). These
are: The presence of a large, nearly trian-
gular, hiatus among the muscles covering
the anterior chamber of the swimbladder
VOLUME 113, NUMBER 1
LY.
Fig. 6. Kolpotocheirodon theloura, new species, paratype, MCP 11161, c&s male, SL 25.9 mm. Anterior
portion of anal fin bearing hooks. Figure shows anal-fin hooks positioned along posterolateral border of anal-
fin rays, bent anteriorly over lateral surface of anal-fin ray to which it is attached, and distal tip pointing
anteriorly.
between the first and second pleural ribs
(pseudotympanum—Figs. 1—3). This hiatus
is limited dorsally by the lateralis superfi-
cialis muscle, posteriorly by a naked ante-
rior face of the second pleural rib, postero-
ventrally by the obliquus inferioris muscle,
and antero-ventrally by the obliquus super-
ioris muscle. See also Malabarba (1998:
200-201, figs. 2B and 3A) and Weitzman
& Malabarba (1999:7 and 16, figs. 5, 6, 16
and 17). The humeral spot is absent (Figs.
1—3). The teeth are pedunculated, largely
expanded and compressed on their distal
borders (Fig. 7). An unique regular, single
tooth row is present on the premaxilla. The
teeth of this row are perfectly aligned and
similar in shape and cusp number (Fig. 7).
Kolpotocheirodon theloura is included
among the members of the cheirodontine
tribe Compsurini (Malabarba, Weitzman, &
Burns in Malabarba, 1998) because it has
the following synapomorphies of that tribe:
The species is inseminating. The anal-fin
hooks are positioned along the posterolat-
eral border of the anal-fin rays, but are bent
more or less anteriorly over the lateral sur-
face of the anal-fin ray to which each is
attached. The distal tip of each anal-fin
hook is directed towards the anterior border
of the anal-fin ray to which it is attached
(Fig. 6), instead of pointing posteriorly or
dorsally as in most other characids that
have anal-fin hooks. Hooks are present on
the dorsal surface of some of the caudal-fin
rays and inclined towards the caudal-fin
base (Fig. 4). Hooks are distributed along
the distal lengths of the principal caudal-fin
rays 11 to 14 (Fig. 4). The anal fin is more
strongly pigmented along the distal portion
of the branched rays (Fig. 1). Kolpotochei-
rodon theloura was placed as the most bas-
al species (identified as Species A) of the
Compsurini in the parsimony analysis of
Malabarba (1998). We herein further dis-
cuss and describe the characters of K. thel-
oura in order to better hypothesize its re-
lationships to other compsurin species.
The presence of insemination, where
278
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs 7.
maxilla and premaxilla and teeth, internal view.
sperm are introduced into the ovary (noted
above), is so far found in the species of the
Glandulocaudinae (Burns et al. 1995), in
the species of the tribe Compsurini of the
Cheirodontinae (Burns et al. 1997; Mala-
barba, Weitzman, & Burns in Malabarba
1998), and species of uncertain relation-
ships including Monotocheirodon pearsoni
Eigenmann in Pearson (1924), Brittan-
ichthys axelrodi Géry (1965), “‘“Cheirodon”’
ortegai Vari & Géry (1980) and “‘Brycon-
americus’’ pectinatus Vari & Siebert
(1990), (Weitzman & Malabarba 1998,
Burns et al. 1999). The absence of cheiro-
dontine synapomorphies among the taxa
Kolpotocheirodon theloura, new species, paratype, MCP 11161, c&s male, SL 25.9 mm. Right
just listed above and the hypothesis that the
Compsurini is a derived branch arising
from externally fertilizing cheirodontines
(see cladogram, fig. 1 of Malabarba 1998)
suggests a hypothesis that insemination in
the compsurin cheirodontines arose from
externally fertilizing ancestral cheirodonti-
ne stock, and that it probably arose inde-
pendently of the Glandulocaudinae and of
the other taxa listed above. At least all chei-
rodontines, including the compsurins, lack
the following synapomorphy of the glan-
dulocaudines used in part to diagnose that
subfamily by Weitzman & Menezes (1998):
presence of a sperm storage area in the tes-
Fig. 8.
illustrates the pelvic-fin rays and hooks in the left pelvic fin, anterior to left. Ventral view.
Kolpotocheirodon theloura, new species, paratype, MCP 11161, c&s male, SL 25.9 mm. Figure
VOLUME 113, NUMBER |
tes. Although rather tentative because not
all glandulocaudine taxa have yet been
sampled, the presence of an elongate cyto-
plasmic collar binding the flagellum to the
nucleus at some stage of spermiogenesis or
in mature sperm cells in glandulocaudines
(see Burns et al. 1998) is absent in at least
one compsurin species, Macropsobrycon
uruguayanae.
The presence of elongate sperm nuclei
was independently pointed out by Weitz-
man & Menezes (1998) as a synapomorphy
for the Glandulocaudinae, and by Malabar-
ba et al. in Malabarba (1998) as a synapo-
morphy for a subclade of the Compsurini
(excluding K. theloura). This requires some
explanation. Sperm with spherical or slight-
ly oval nuclei are often referred to as
‘“‘aquasperm’”’ and considered to be the ple-
siomorphic sperm type for teleosts (Jamie-
son 1991). Elongation of sperm nuclei is a
derived feature and seems to be related to,
but not necessarily needed for, sperm trans-
fer to and storage within ovaries (Jamieson
1987). Such a modification is found in the
two inseminating characiform groups listed
above and in several inseminating or inter-
nally fertilizing teleosts. Although the
sperm cells in these groups have undergone
changes in shape resulting in sperm cells
different from the spherical or slightly oval
nuclei of “‘aquasperm,’’ these changes are
not necessarily homologous. Although we
still lack information on the ultrastructural
morphology of the sperm cells of most
compsurins and several glandulocaudines,
the evidence so far noted in Burns et al.
(1998) indicates the elongated sperm cells
are derived in different ways in the comp-
surin Macropsobrycon uruguayanae and in
the glandulocaudines Diapoma speculifer-
um Cope (1894), Diapoma sp., Corynopo-
ma riisei Gill (1858), Pseudocorynopoma
doriae Perugia (1891), Mimagoniates bar-
beri Regan (1907), and M. microlepis
(Steindachner, 1876). Thus, elongation of
the sperm nuclei in glandulocaudines and
elongation of the sperm nuclei in at least
part of the compsurins appear to have been
279
acquired in different ways and are non-ho-
mologous, and thus likely constitute differ-
ent synapomorphies for each clade. How-
ever, much of the ultrastructural evidence
from sperm cell morphology in the species
of both clades remains to be discovered and
described.
The anal-fin hook arrangement of the
Compsurini, including Kolpotocheirodon
theloura, is also hypothesized to constitute
a synapomorphy. Compsurin cheirodon-
tines, except for Acinocheirodon melano-
gramma, have hooks inserted along the
posterolateral border of the anal-fin rays
and bent anteriorly over the lateral surface
of the anal-fin ray where each is attached
(Fig. 6). Thus, instead of pointing posteri-
orly and dorsally as in most characids, the
distal tip of the anal-fin hooks are directed
anteriorly. Non-compsurin cheirodontines
have retrorse hooks positioned along the
posterolateral border of the anal-fin rays,
and directed posteriorly and towards the
anal-fin base. Some glandulocaudines such
as the xenurobryconins have the anal-fin
hooks recurved and pointing dorsoanterior-
ly, but in most cases these differ in shape
from those of the compsurins, do not bend
anteriorly over the lateral surface of the
anal-fin ray, and do not pass across the
point where it is attached (see Weitzman &
Fink 1985:figs. 38-43; Weitzman et al.
1994:fig. 11; and Weitzman & Ortega,
1995:fig. 4). Among xenurobryconins, Ar-
gopleura chocoensis Eigenmann has modi-
fied large anal-fin hooks, anterodorsally and
laterally directed, on rays 6—7 to 12-14, re-
sembling those of the compsurins (see
Weitzman & Fink 1985:fig. 38). However,
these hooks in A. chocoensis are part of a
complex pattern of hook development (see
Weitzman & Fink 1985:29, apomorphy 7)
absent in compsurin species. In any case,
parsimony analysis also indicates that the
condition in the Xenurobryconini and the
Compsurini are non-homologous.
The presence of hooks on the dorsal sur-
face of some of the caudal-fin rays (char-
acter 7) is also found among inseminating
280
glandulocaudines (see Menezes & Weitz-
man 1990:fig. 5, Mimagoniates microlepis
and fig. 24, Mimagoniates rheocharis Me-
nezes & Weitzman (1990); Weitzman &
Fink 1985:fig. 11, Xenurobrycon macropus
Myers & Miranda-Ribeiro (1945), fig. 12,
Xenurobrycon heterodon Weitzman & Fink
(1985), fig. 16, Corynopoma riisei, fig. 17,
Gephyrocharax atricaudatus (Meek & Hil-
debrand 1912), fig. 19, Mimagoniates mi-
crolepis; Weitzman et al., 1994:fig. 5,
Ptychocharax rhyacophila Weitzman et al.
(1994), in the incertae sedis species Brit-
tanichthys axelrodi (see Malabarba &
Weitzman 1999:figs. 9-10) and “‘Cheiro-
don’”’ ortegai. The possible homology of the
caudal fin modifications of the inseminating
compsurin cheirodontine Acinocheirodon
melanogramma with two Mimagoniates
Regan (1907) species and with B. axelrodi
were extensively discussed in Malabarba &
Weitzman (1999) and rejected. Homology
of the caudal-fin hooks in K. theloura with
all non-compsurin taxa listed above is also
rejected because these inseminating chara-
cids lack the synapomorphies diagnosing
the Cheirodontinae.
The putative caudal gland of Kolpoto-
cheirodon theloura does not have gross
similarities to those of the inseminating
glandulocaudines, except possibly Landon-
ia latidens Eigenmann & Henn in Eigen-
mann et al. (1914) of the tribe Landonini
that also has folds of tissue in the caudal-
fin region. Although a detailed comparison
of the histological nature of these species
needs to be done, it is very unlikely the
folds in the Compsurini and Landonini are
homologous in a parsimony analysis. The
caudal organ of K. theloura also does not
resemble those found in Compsura and
Saccoderma. Species of these genera have
derived scales associated with their caudal
organs. Instead the caudal organ of K. thel-
oura is formed by derived dermal flaps
found along the proximal length of the thir-
teenth or fourteenth to nineteenth caudal-fin
rays. Although not sharing a tabular or pa-
pillose margin, the caudal glands of other
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
inseminating compsurins are also partially
formed by derived hypertrophied dermal
flaps along the ventral lobe caudal-fin rays,
as described by Malabarba & Weitzman
(1999) along the thirteenth through fifteenth
caudal-fin rays of Acinocheirodon melano-
gramma, and fifteenth or sixteenth to nine-
teenth caudal-fin rays in the Saccoderma
species and Compsura heterura Eigen-
mann. The caudal glands of compsurin spe-
cies, although formed in different ways, are
always derived from modified dermal flaps
along the ventral lobe caudal-fin rays. This
seems to reinforce the monophyly of the
group and the hypothesized homology or
common origin of insemination among
compsurin cheirodontines.
Males of Kolpotocheirodon theloura
have modified twelfth to the fourteenth or
fifteenth principal caudal-fin rays. These
rays bear 4—6 retrorse hooks on each side
in a row along their dorsal divisions (Fig.
4). Among the Compsurini, Malabarba &
Weitzman (1999) listed and described the
presence of hooks in Acinocheirodon me-
lanogramma (caudal-fin rays 13-14, rarely
on ray 15), Saccoderma hastata Eigenmann
(1915) (caudal-fin rays 13-18), ‘‘Odonto-
stilbe”’ dialeptura (caudal-fin rays 12—16),
and in Macropsobrycon uruguayanae (cau-
dal-fin rays 12—14, plus several spinelets
along the proximal half of the caudal-fin
rays 14 to 18). Although these hooks are
present on the ventral lobe of the caudal fin
in all these species, we noted (Malabarba &
Weitzman 1999) that they do not all occur
on the same caudal-fin rays in all species
and are of different shapes. Additionally,
not all compsurin species have caudal-fin
hooks, as for example Compsura heterura,
Compsura gorgonae Evermann & Golds-
borough and ‘‘Odontostilbe’’ mitoptera. We
also pointed out that the use of the presence
of caudal-fin hooks in diagnosing a clade
among the Compsurini species would be
parsimoniously incongruent with other de-
rived characters, as for example the pres-
ence of derived pouch scales in the ventral
lobe of the caudal fin, found in some hook-
VOLUME 113, NUMBER |!
bearing Saccoderma species and some non-
hook-bearing species such as Compsura
heterura and C. gorgonae. A parsimony
analysis of character distribution indicates
the presence of hooks as a synapomorphy
for the compsurin cheirodontines, and its
lack a secondary reversal in some of its spe-
cies (Malabarba et al. in Malabarba 1998).
Kolpotocheirodon theloura was placed as
the most basal species among the Comp-
surini, in the phylogeny proposed by Mal-
abarba et al. (in Malabarba 1998). Among
the characters that placed the new species
as the most basal compsurin is the presence
of aquasperm (see Burns et al. 1997:434,
fig. 1A), as found in externally fertilizing
cheirodontines. In other words, the presence
of derived elongate sperm nuclei (see Burns
et al. 1997:434, table I) in the remaining
inseminating compsurins represents a syn-
apomorphy for these derived fishes.
Another character that seems to corrob-
orate a basal position for Kolpotocheirodon
theloura is the lack of enlarged caudal-
pouch scales as a possible pumping mech-
anism in the ventral lobe of the caudal fin
associated with a caudal gland (see Weitz-
man & Fink 1985:96—-99; Malabarba &
Weitzman 1999). Compsura heterura, C.
gorgonae, and Saccoderma species have
the posterior ultimate scale of the scale row
immediately ventral to the lateral-line scale
row enlarged, connected dorsally to liga-
ments arising from the twelfth and _thir-
teenth principal caudal-fin rays, and ven-
trally to a skin flap connected to the nine-
teenth principal caudal-fin ray, forming a
pouch with a posterior opening that may
serve aS a pumping mechanism. Acino-
cheirodon melanogramma does not have
any pumping mechanism involving caudal-
fin scales. Instead it has two large skin flaps
between the thirteenth and fifteenth princi-
pal caudal-fin rays (Malabarba & Weitzman
1999:fig. 6), that occur just ventral to and
surrounding the hypertrophied soft tissue
found around the hooks of the thirteenth
and fourteenth fin rays. Malabarba &
Weitzman (1999) suggested that these flaps
281
might also be part of a mechanism for in-
creasing the rate of pheromone dissemina-
tion in the surrounding water during court-
ship. We found no putative active pumping
mechanism in Kolpotocheirodon theloura
that might be related to an increase in pher-
omone dissemination in water during court-
ship. The papillose margins of the dermal
flaps of K. theloura seem to be a passive
strategy for increasing pheromone dissem-
ination.
Acknowledgments
Financial aid for travel for museum study
and field work in Brazil were supplied to
LRM by CAPES and Conselho Nacional de
Desenvolvimento Cientifico e Tecnolégico
(CNPq-Proc. 451459/96-2). José Lima de
Figueiredo and Osvaldo Oyakawa (MZUSP)
loaned specimens in their care or provided
cataloging services. Lisa Palmer (USNM)
and José EF P. Silva provided cataloging and
other technical services. John R. Burns pro-
vided useful insights through histological
and TEM analyses. This paper was im-
proved by the comments of José Lima de
Figueiredo in the first version of the descrip-
tion of this species, and in more recent ver-
sions by Richard Vari.
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):284—290. 2000.
Calls and calling behavior of the frog Leptodactylus natalensis
(Amphibia: Anura: Leptodactylidae)
W. Ronald Heyer and Celso Morato de Carvalho
(WRH) Department of Vertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, DC 20560-0107, U.S.A.; (CMC) Departamento de Biologia,
Universidade Federal de Sergipe, Jardim Rosa Elze, s/n, Sao Crist6vao, SE 49100-000, Brazil
Abstract.—Leptodactylus natalensis has a prolonged calling season, typically
from May to September. Individuals occur in and call from the most forested
patch of habitat on the campus of the Universidade Federal de Sergipe. Two
kinds of calls were recorded and analyzed—advertisement and chirp calls. The
function of the chirp call is unknown. The advertisement call has two carrier
frequency bands, known to date for only two other species of Leptodactylus.
The two carrier bands may correspond to different frequency tuning sensitiv-
ities in males and females, as females have slightly larger tympani than males,
corresponding with their larger overall size.
Published natural history information for
the frog Leptodactylus natalensis Lutz,
1930, consists of comments on calling and
microhabitat made by Lutz (1930) in his
original description of the species and a re-
cent study of some reproductive aspects, in-
cluding the first description of the tadpole
for the species (Oliveira & Lirio Junior
1999). Calls of Leptodactylus natalensis
have not been analyzed previously (Heyer
1994). One of us (CMC) had the opportu-
nity to observe calling behavior and record-
ed the calls of L. natalensis. The purpose
of this paper is to report these observations
and describe the calls.
Methods and Materials
Observational data and recordings of
Leptodactylus natalensis were made by
CMC on the campus of the Universidade
Federal de Sergipe, SAo Crist6vao, Sergipe,
Brazil. Incidental information on calls at
Santo Amaro das Brotas, Sergipe, was also
obtained by CMC.
Recordings were made from two individ-
uals. Five recordings (USNM tape 323, cut
1) were made from specimen MZUSP
89945 (MZUSP = Museu de Zoologia,
Universidade de Sao Paulo collection),
SVL (snout—vent length) 33.9 mm, on 22
June 1998, at an air temperature of 25°C,
water temperature 24°C, the last recording
completed at. 1955 h. One recording
(USNM tape 323, cut 2) was made from
specimen MZUSP 89947, SVL 32.4 mm,
also on 22 June 1998, air 25°C, water 24°C,
the recording completed at 2030 h. It was
not raining when these recordings were
made, but a brief heavy rain occurred at
2100 h.
Calls were recorded with a Sony TM-
5000 cassette recorder with a M 44 N(C)
Beyer Dynamic Soundstar II microphone.
Terminology follows that of Heyer et al.
(1990), with the following clarifications.
Carrier frequency is the frequency broad-
cast by the calling frog to the environment.
Often, the carrier frequency embraces a
range of frequencies loud enough to be
heard by the intended receiver (male or fe-
male). The dominant frequency as used
here is the single frequency value (or very
narrow range of frequencies) that has the
most sound energy in the call.
Recordings were analyzed using ‘‘Ca-
VOLUME 113, NUMBER 1
nary 2.1”’ software (Charif et al. 1995). Call
rates and durations were measured from
wave form displays, beginning and ending
carrier frequencies from audiospectrogram
displays, and dominant frequencies from
spectral displays. Although the initiation of
calls is clear on the wave form displays, the
ends of the calls are not very distinct, per-
haps due to the level of background noise
and possible microphone ringing.
Advertisement call rates were based on
six calling sequences ranging from 7.8 to
15.2 sec from each of six recordings. Fre-
quency data were taken from 10 consecu-
tive calls from each of the six recordings;
call duration and pulse data were taken on
the same 10 calls for recordings 2 and 5 of
MZUSP 89945 and for MZUSP 89947.
Chirp call data are from the six chirps
loud enough to analyze in the second re-
cording of MZUSP 89945.
Behavior and Calling Site
Leptodactylus natalensis has a prolonged
calling season coinciding with the rainy
season, typically from May to September
(also see Oliveira & Lirio Junior 1999).
Calling starts around 1600 h. Calling activ-
ity is most intense around 1900 and contin-
ues until 2300 or 2400 h. However, on rainy
days, especially at the beginning of the
rainy season, males can be heard calling at
0800 h, and continuing throughout the day
and night. Males do not call during hot and
sunny days during the rainy season.
The general site the frogs occupy on the
campus is a small patch (100 <X 400 m) of
second growth (capoeira) on the right bank
of the Rio Poxim, a seasonally partially
flooded area (varzea) with many ponds. The
site has 5—6 m high trees, many young palm
trees, a thick grass ground cover, shallow
litter, and white sand. The Leptodactylus
natalensis were only heard calling from the
capoeira—never from the left bank of the
Rio Poxim, which has more artificial open
areas and patches of mangrove swamps.
MZUSP 89945 was calling very close to
285
the foot of a young palm tree surrounded
by a pond with leaves in the water. The frog
sat in the water, half submerged. A second
male (MZUSP 89946) was about a half me-
ter from the calling male. This second male
was in a small burrow, with half the body
exposed and uttered some “‘clicks”’ but no
other calls. The “‘clicks’’ did not record ad-
equately enough to analyze.
MZUSP 89947 was near the foot of a
small tree, at the edge of a little channel of
rain water, hidden in grass.
Leptodactylus natalensis infrequently
jump when disturbed. Usually they take two
or three steps backward and quietly hide
under a leaf or other ground cover.
Advertisement Call
Calls (Fig. 1) consist of single notes giv-
en frequently when actively calling. Call
rates vary from 3.4—4.1 per sec for MZUSP
89945 and at a rate of 3.0 per sec for
MZUSP 89947. The call sequence with the
least background noise is the fifth recording
sequence for MZUSP 89945. In that se-
quence, each call (=a single note) has a du-
ration of 0.06—0.07 (mode = 0.07) sec and
consists of 5—7 pulses, modally 7. The call
is partially pulsed (=incompletely ampli-
tude modulated). In two other recordings
with greater background noise, it appears
that the terminal pulses are indistinguish-
able from the background noise in the wave
forms. Three to five partial pulses (mode =
4) can be distinguished in calls from the
second recording of MZUSP 89945, with
an average duration of 0.04 sec. Two to
three partial pulses (mode = 3) are distin-
guishable in the recording from MZUSP
89947, with an average duration of 0.02
sec. The calls are frequency modulated with
extremely fast rise times (difficult to see be-
cause of time scale in Fig. 1, more visible
in the two advertisement calls shown in Fig.
3). Beginning frequencies of the carrier
(=fundamental) frequency range from av-
erages of 550—600 Hz with the highest fre-
quencies ranging from averages of 1370-—
286 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
“ni Wernher
50 ms
Sa. Saar ls
N
ed
i
iP)
c=
‘e)
eas
v
e e
Time in seconds
Fig. 1. Wave form (above) and audiospectrogram (below) of the advertisement call of Leptodactylus natal-
ensis, USNM tape 323, cut 1, fifth recording. The wave form is of the first call shown in the audiospectrogram.
1830 Hz for MZUSP 89945. The average
beginning dominant frequency for MZUSP
89947 is 650 Hz, with an average maxi-
mum of 1600 Hz. The dominant frequen-
cies most often occur in two peaks in the
calls of MZUSP 89945. For this frog, the
first recording series usually has a dominant
frequency of 760 Hz, with either a co-dom-
inant or quieter frequency at 1050 Hz; the
second recording has the dominant frequen-
cy at 770 Hz, with a secondary dominant
at 1020 Hz; the third recording has the
dominant frequency at 760 Hz, with a sec-
ondary dominant at 1040 Hz; the fourth re-
cording has the most variation with either
780 and 1030 Hz being primary or second-
ary in loudness, 860 and 1030 Hz being
primary or secondary in loudness, 1030 Hz
loudest with a secondary dominant at 880
Hz, or a single dominant frequency at 1030
Hz; calls in the fifth recording have the
dominant frequency at 1030—1040 Hz, with
a secondary dominant at 760—770 Hz (Fig.
2). The calls of MZUSP 89947 have either
a dominant frequency at 1030 Hz, or also
with a secondary dominant at 875 Hz. The
partial pulsing of the calls appears to be the
cause of the sidebands weakly visible in the
audiospectrograms (Fig. 1). Calls of
MZUSP 89945 range from having no visi-
ble harmonics, a weak second harmonic, or
intermediate strength second and third har-
monics (Fig. 2). The calls of MZUSP
89947 have a weak second harmonic.
Chirp Call
Calls (Fig. 3) consist of either 1 or 2
notes. The loudest chirp calls are louder
than the advertisement calls. In the two note
VOLUME 113, NUMBER 1
dB
287
| | 1980-2070
\ as
VV \ 2600 2930
oe Ch ow ee el aoe
O 1 2 3 + 5
e
Kilohertz
Fig. 2. Power spectrum of advertisement call of Leptodactylus natalensis, USNM recording 323, cut | (of
second call in audiospectrogram of Fig. 1).
calls, the first note is shorter than the sec-
ond (0.04 and 0.13 with an internote inter-
val of 0.10 sec; 0.05 and 0.08 with an in-
ternote interval of 0.16 sec; 0.04 and 0.13
with an internote interval of 0.14 sec). The
three single note chirp calls recorded range
from 0.10—0.14 sec duration. From 3-10
partial pulses can be discerned in the wave
forms. The calls are frequency modulated,
at a noticeably slower rise time than the ad-
vertisement calls. The lowest carrier (=fun-
damental) frequency ranges from 530—650
Hz; the highest carrier frequency ranges
from 990-1200 Hz. There is a suggestion
of the carrier frequency briefly falling at the
end of some calls, back to around 900—950
Hz. The dominant frequencies vary from
690-860 Hz. There are weak to strong sec-
ond, second and third, or second, third, and
fourth harmonics.
Discussion
The function of the chirp calls is un-
known. Other members of the L. melano-
notus group have these calls as well, which
usually initiate calling bouts. In the case of
the L. natalensis recording, the chirp calls
were uttered after there was a slowing down
in the rate of advertisement calls. The chirp
calls are much more variable than the ad-
vertisement calls.
Many species of Leptodactylus have fre-
quency modulated calls that result in broad-
casting their calls over a range of frequen-
cies. Most of these calls have a single (or
narrow range) dominant frequency, how-
ever. Leptodactylus natalensis is unusual in
that there are two peaks of loudness. Only
L. melanonotus and podicipinus within the
L. melanonotus group share this condition
(of those for which the calls are known, Ta-
ble 1). We hypothesize that these separate
dominant frequencies correspond to differ-
ent tuning curves in males and females, as
demonstrated in Eleutherodactylus coqui
(Narins & Capranica, 1976), although no
experimental data are available for L. na-
talensis. Morphological data do not contra-
dict this hypothesis. Other things being
equal, a larger tympanum will be more sen-
sitive to lower frequencies than a smaller
tympanum and vice versa. Morphological
b>
Kilohertz
NO
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Time in seconds
Bigs:
Wave form (above) and audiospectrogram (below) of the chirp call of Leptodactylus natalensis,
USNM tape 323, cut 1. In the audiospectrogram, a single-note chirp call is followed by two advertisement
calls.
data are available from two localities rep-
resented by several males and females. The
data indicate that the tympanum size does
differ between males and females (Table 2).
It would seem from the variability of calls
in the fourth recording of MZUSP 89945
that an individual frog has control over
whether more energy is broadcast in the
lower or higher dominant frequency.
Sexual difference in tympanum size is the
expected condition in most frogs, as most
frog species are sexually dimorphic, with the
females being larger. Broadcasting of higher
and lower dominant frequencies, as seen in
L. natalensis, is one solution to matching
different receptor sensitivities to the adver-
tisement call. The frequency modulation ob-
served in other Leptodactylus species, such
as L. fuscus, is another solution.
Our observations apparently conflict with
those of Adolfo Lutz regarding the call of
L. natalensis. In the original description of
the species, Lutz observed (1930:27, En-
glish version of text): ““This frog has a call
with the strength of a whistle but sounding
more like the voice of a cricket. It seems to
have also another sound, heard during cop-
ulation, like a soft clucking, sometimes re-
peated by other males, so as to form a gen-
eral concert. The specimens were caught
near to the water but generally well hid-
den.’? CMC has not heard cricket-like calls
from L. natalensis at either the campus at
Sao Crist6vao or Santo Amaro das Brotas,
VOLUME 113, NUMBER 1
289
Table 1.—Advertisement call characteristics of members of the Leptodactylus melanonotus species group.
Data taken from Heyer (1994), Heyer et al. (1996), and for L. melanonotus, from USNM tape 83, cut 1. Note
that Leptodactylus diedrus is no longer considered a member of the L. melanonotus species group (Heyer, 1998).
The duration and pulse data for L. natalensis are from the recording with the least background noise.
Call duration Number partial
Carrier Dominant
Species in sec pulses per call frequency range frequency range
L. griseigularis 0.01 l 1380-3060 2770
L. leptodactyloides 0.02—0.04 1 or 3-5 650-1600 1 100—1300
L. melanonotus 0.07—0.08 4-6 820-2620 1320-1380 or 2330
L. natalensis 0.06—0.07 5-7 550-1830 760-880 or 1020—1040
L. nesiotus 0.03 4-5 1500—2000 1800—2000
L. pallidirostris 0.03-—0.05 1 or 2-5 1500-3500 3000-3500
L. petersii A 0.04—0.05 3-4 700-1200 750-800
L. petersii B 0.03—0.05 1 or 2-4 800—1600 or 1800—2800 N.A.
L. podicipinus 0.02—0.04 3-7 1000—3500 1000—1200 or 3300-3500
L. sabanensis 0.04—0.06 very weak 900-2300 1400—1800
L. silvanimbus 0.15 about 20 420-1920 510
L. validus 0.03—0.06 1 or 2-6 1300-3500 2300-3500
but the call we describe as the advertise- Acknowledgments
ment call is a soft clucking call.
Advertisement calls in the L. melanono-
tus group are often not as species-specific
as in other Leptodactylus, perhaps because
the calls have a strong point-location ori-
entation function overriding (or in addition
to) a species-coding function (Table 1 and
Heyer 1994:106—107). Leptodactylus natal-
ensis has a disjunct geographic distribution
relative to all other members of the L. me-
lanonotus group. The species group mem-
ber that most closely approaches the distri-
bution of L. natalensis is L. podicipinus
(compare Figs. 39 and 43 in Heyer 1994).
Interestingly, as pointed out above, L. pod-
icipinus is only one of two other known
members of the L. melanonotus group be-
side L. natalensis to have two distinct loud-
est broadcast peaks in the dominant fre-
quency band.
This research was partially supported by
the Neotropical Lowlands Research Pro-
gram, National Museum of Natural History,
Dr. Richard P. Vari, Principal Investigator
(WRH) and Departamento de Biologia,
Universidade Federal de Sergipe (CMC).
We thank these institutions for support and
Dr. A. Stanley Rand for reviewing drafts of
this paper for us.
Literature Cited
Charif, R. A., S. Mitchell, & C. W. Clark. 1995. Ca-
nary 1.2 User’s Manual. Ithaca, New York, Cor-
nell Laboratory of Ornithology.
Heyer, W. R. 1994. Variation within the Leptodactylus
podicipinus-wagneri complex of frogs (Am-
phibia: Leptodactylidae).—Smithsonian Contri-
butions to Zoology 546:1—124.
1998. The relationships of Leptodactylus
Table 2.—Snout-vent length and tympanum diameter measurements for male and female Leptodactylus na-
talensis from two localities in the State of Sergipe, Brazil.
Snout—vent length Tympanum diameter
Locality n Sex Min. Max. Mean Min. Max. Mean
Areia Branca 8 fe) 28.7 36.5 33.1 Zz 32 2.8
Areia Branca 9 Q 33.9 42.6 38.8 Pink 3.4 3.0
Santo Amaro das Brotas iT 3 30.6 34.6 325 2.4 ZF 2.6
Santo Amaro das Brotas 14 2 33.1 37.0 S| Ps 2.9 Og |
290 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
diedrus (Anura, Leptodactylidae).—Alytes 16: ras alliadas——Memorias do Instituto Oswaldo
1294. Cruz 23:1-—20 (Portuguese), 21—34 (English),
, R. de SA, J. R. McCranie, & L. D. Wilson. +5 plates.
Narins, P. M., & R. R. Capranica. 1976. Sexual dif-
ferences in the auditory system of the tree frog
Eleutherodactylus coqui.—Science 192:378-
380.
Oliveira, EF FE, & G. P. Lirio Junior. 1999. Diversidade
e aspectos reprodutivos de uma comunidade de
1996. Leptodactylus silvanimbus (Amphibia:
Anura: Leptodactylidae): natural history notes,
advertisement call, and relationships.—Herpe-
tological Natural History 4:169-174.
, A. St Rand, C, A: G, Cruz, OF} Peixoto, &
Caples Nelson. fe Frogs of Boracéia.—Ar- anfibios anuros de uma regiao do rio Poxim,
quivos de Zoologia 31:231—410. Sergipe (Amphibia, Anura).—Publicag6es
Lutz, A. 1930. Segunda memoria sobre especies bras- Avulsas do Centro Académico Livre de Biolo-
ileiras do genero Leptodactylus, incluindo out- gia, Universidade Federal de Sergipe (in press).
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):291-297. 2000.
A new species of venomous coral snake (Serpentes: Elapidae) from
high desert in Puebla, Mexico
Jonathan A. Campbell
Department of Biology, The University of Texas at Arlington,
Arlington, Texas 76019, U.S.A.
Abstract.—A new species of coral snake (Micrurus pachecogili) is described
from southern Puebla, Mexico. Black body rings in this snake are monadal and
the species is allopatric from all other coral snakes except perhaps M. laticol-
laris, a species in which black body rings are in triads. The new species is
characterized by distinctive, well developed yellow rings, relatively short red
rings, and a relatively large number of ventral scutes in comparison with M.
nebularis.
Resumen.—Se describe una nueva especie de serpiente coral (Micrurus pa-
checogili) del sur de Puebla, México. En esta serpiente, los anillos corporales
negros son monodales; la nueva especie es alopatrica de otras serpientes de
coral, tal vez con excepcién de Micrurus laticollaris, una especie en la que los
anillos negros se disponen en triadas. La nueva especie se caracteriza por po-
seer anillos amarillos bien desarrollados, anillos rojos relativamente cortos y
un numero grande de escamas ventrales en comparacion con M. nebularis.
Species of coral snakes are widespread,
if not always conspicuous, components of
New World herpetofaunal assemblages.
They occur throughout most of the Neo-
tropics and reach into both temperate North
and South America. These snakes are often
associated with mesic conditions and the
majority of species occur in rainforest or
cloudforest habitats. Coral snakes appear to
be delicate and desiccate quickly when ex-
posed to direct sunlight. Thus, most species
are nocturnal or are active only during early
morning or later afternoon, especially on
rainy nights or overcast days. However, a
fair number of coral snakes have become
adapted to subhumid habitats, occurring in
desert, thorn scrub, or tropical deciduous
forest from the Sonoran Desert of the south-
western U.S.A. southward on the Pacific
coast of Mexico and Central America and
in various regions in South America. In Pa-
cific Mexico, tropical deciduous forest is in-
habited by Micurus browni, M. distans, M.
ephippifer, M. bogerti, M. laticollaris, M.
proximans, and Micruroides euryxanthus,
with the latter species also extending into
the Sonoran Desert (Campbell & Lamar
1939).
The arid lands of southern Puebla are sit-
uated in the northern part of the Mesa del Sur
physiographic region and are just south of the
Mesa Central (West 1964, fig. 3). Heretofore,
this area was reported to harbor only a single
species of coral snake, M. laticollaris (Camp-
bell & Lamar 1989, Roze 1996), easily dis-
tinguishable from other Mexican coral snakes
in being one of two species with black body
rings in triads (the other triadal species being
M. elegans of mesic forests on the eastern
versant). Some years ago, I traversed on mul-
tiple occasions the countryside surrounding
Zapotitlan Salinas in southern Puebla, during
which time I assembled small collections of
amphibians and reptiles. Among the material
in these collections are two specimens of a
species of coral snake that cannot be associ-
ated with any species known from Mexico.
292
Materials and Methods
Descriptions of characters and terminol-
ogy of scales in the diagnosis and descrip-
tion of this new taxon follow Campbell &
Lamar (1989) and Roze (1996). The sex of
individuals was confirmed by checking for
the presence of hemipenes by making a
small midventral incision on the proximal
section of the tail. Head measurements were
taken to the nearest 0.1 mm using digital
calipers held under a dissecting scope and
the snout—vent and total length was taken
to the nearest 1.0 mm using a meter stick.
Micrurus pachecogili, new species
(Figs. 1-3)
Holotype.—An adult male (Figs. 1-2),
UTA R-12546 (original number JAC 9752),
from 5.6 km SSW Zapotitlan Salinas, 1494
m, Puebla, Mexico, collected in November
1983 by one of the children of E. Pacheco-
Gil. This locality is in high tropical arid for-
est at 18°18'N, 97°31'W (Fig. 3).
Paratype.—A subadult male, UTA R-
17145, from the type-locality, collected in
December 1985.
Diagnosis.—Micrurus pachecogili dif-
fers most notably from other species of Mi-
curus in central Mexico in being tricolored
and having black body rings in monads
(i.e., a body pattern of red-yellow-black-
yellow-red), in having 220—223 ventrals in
males, in possessing a wide pale parietal
ring that includes all of the parietals and
portions of adjacent scales, and in having
non-melanized red body rings that are about
equal to or shorter than black body rings.
Indeed, these four characters, in combina-
tion, distinguish M. pachecogili from all
other venomous Mexican coral snakes.
Micrurus bernadi differs in having mostly
red and black body rings, although a yellow
parietal ring and, rarely, narrow yellow edg-
ing may be present along black markings,
black rings are usually reduced to dorsal
spots or saddlelike bands, and males have
198-212 ventrals. Micrurus browni differs
in having a black nuchal ring that covers the
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
posterior portion of the parietals and adult
males have supracloacal keels. In M. b.
browni of Guerrero and Oaxaca, the subspe-
cies that is geographically most proximate to
Zapotitlan, males have 204—218 ventrals, the
black head cap extends to the anterior part
of the parietals and frontal, and the black
nuchal ring covers the ultimate and penulti-
mate supralabials (versus only the posterior
edge of the ultimate supralabial). Micrurus
diastema differs in that the red rings are ei-
ther very long (>5 times wider than the
black rings) or contain scales that are dis-
tinctly black-tipped, the black head cap in-
cludes the anterior portion of the parietals
and/or the black nuchal ring includes the
posterior portion of the parietals. Micrurus
distans of western Mexico differs in having
no more than 17 black body rings, males
have <217 ventrals, and red body rings are
usually at least three times longer than black
rings. Micrurus ephippifer differs in having
a black nuchal ring that covers the parietal
tips, red rings have black-tipped scales,
spots, or large saddlelike bands, and =219
ventrals in males. Micrurus laticollaris has
black body rings arranged in triads (i.e.,
body pattern of red-black-yellow-black-yel-
low-black-red), has 1 + 2 temporals, and
males possess $215 ventrals. Micrurus lim-
batus is bicolored with red and black rings
and <192 ventrals. Overall, the color pattern
of M. pachecogili is most similar to that of
M. nebularis: similarities include red and
black rings of relatively equal length, scales
in red rings not black-tipped or distinctly
spotted, well developed yellow body rings
(1.0—1.5 dorsal scale lengths long in M. ne-
bularis versus 2.0 in M. pachecogili), and
tail with subequal black and yellow rings.
Micururus nebularis differs by having a
black nuchal ring that covers the posterior
ends of the parietals, 203—208 ventrals in
males, and seven black tail bands.
Description of holotype.—A single pre-
nasal and postnasal on each side; nostril sit-
uated mostly in posterior part of prenasal; an
elongate preocular on each side contacting
postnasal; no loreal; anterior scales on head
VOLUME 113, NUMBER 1
and snout, including mental and infralabials,
with numerous tiny tubercles; postoculars 2/
2; temporals 1 + 1 + 2; upper tertiary tem-
poral large, about half size of parietal; su-
pralabials 7/7; infralabials 7/7 (Fig. 1); ven-
trals 223: cloacal scute divided; subcaudals
47, all divided except proximal subcaudals
2-7; dorsal scale rows smooth, in 15 unred-
uced rows; no supracloacal keels.
Dimensions are as follows: head length
15.0 mm, head width 10.9 mm, total length
639, tail length 87 mm (comprising 13.6%
of total).
The black head cap covers the rostral, first
three supralabials, the upper and anterior
edge of Supralabial 4, the anterior edge of
the upper preocular, slightly more than three-
fourths of the supraoculars, and about half
of the frontal; there is no pale spot on snout.
The yellow parietal ring extends from the
black head cap to include all of the parietals,
the primary and secondary temporals, and
anterior part of the tertiary temporals, and
all of the ultimate supralabial except for the
posterior edge (Fig. 1). About half or slight-
ly more of the lingual portion of the mental
and the first three pairs of infralabials are
black; the posterior gular area is immaculate
yellow and continuous with the yellow pa-
rietal ring. The black nuchal ring does not
reach the posterior tips of the parietals and
involves less than half of the first dorsal
scale situated partially between the parietals.
The nuchal ring extends posteriorly along
ime> middorsum ~1/2 + ° 5 + 1/2 dorsal
scales; it is shortened ventrally to cover most
of the first two ventrals and adjacent 1.5
preventrals. There are 24 black body rings,
including the nuchal, with the posteriormost
just anterior to the vent. Throughout most of
the body black rings are about four dorsal
scale lengths long, but the anterior three
rings behind the nuchal ring are slightly lon-
ger, being about 4.5-5.5 scales in length
(Fig. 2). On the belly, most black body rings
are reduced to about three ventrals (one ring
covers only two ventrals and several cover
four), with an additional half ventral (divid-
ed along midventral line) associated with
293
many rings. Black body rings tend to have
vertical edges or, if shortened on the side of
the body, they are reduced by no more than
about 0.5 scale lengths. The yellow rings are
immaculate and relatively wide throughout
the body, being about two dorsal scales long.
The cloacal scute is yellow. Most red rings
are 3—4 scales in length with a few reduced
to as short as one and one-half to two scale
lengths on the posterior half of the body.
Scales in the red rings are mostly immacu-
late but with slightly dusky free margins and
there are a few irregular, inconspicuous
black dots in a few of the red rings. There
are five black tail rings that are one and one-
half to three times the width of the yellow
rings separating them; the tip of the tail is
yellow.
Variation.—The paratype (UTA R-
17145) is a subadult male, 327 mm in TL,
with a tail length of 36, comprising 11.0%
of the total. It agrees with the holotype in
most respects of scutellation and pattern,
but has 220 ventrals, 43 subcaudals (sub-
caudals 2—3 not divided); a black nuchal
ring involving only the first ventral plus ad-
jacent two and one-half preventrals; 27
black body rings, most of which are about
four dorsal scales in length throughout the
body; red body rings which are two to three
dorsal scales in length throughout the body;
and 6 black tail rings, exclusive of the black
tail tip, which are 1.5—2 times longer than
the yellow rings separating them.
Etymology.—The species name is a noun
in the genitive case, formed in honor of
Emiglio Pacheco Gil, a good friend and
longtime resident of the Zapotitlan Valley.
He, his wife, and about twelve children (I
never knew the total number and am not
sure he did either) always were gracious
and hospitable hosts during my visits, most
willing to share whatever meager provi-
sions they had available. The country
around Zapotitlan is harsh and Emiglio sup-
plemented his income by working in the
onyx mines which abound in the region. He
was killed in a mining accident in 1982.
Remarks.—The morphology of coral
fl
|
| &
VOLUME 113, NUMBER 1
Fig. 2.
snakes is extremely conservative (Slowin-
ski 1995) and the numbers and arrangement
of most head scales, rows of dorsal scales,
and even skull and dentition is remarkably
consistent, with minor exceptions, through-
out Micrurus (Campbell & Lamar 1989,
Roze 1996). Perhaps, the most dramatic ad-
aptation within the genus is the conspicuous
elongation of the body apparent in certain
coral snakes including the genus Leptomi-
crurus and various species of Micrurus in
Lower Central America and northern South
America. Many species of coral snakes are
diagnosed primarily on the basis of color
pattern and this has proven to be a generally
reliable method for distinguishing most spe-
cies, although a few species are notoriously
variable (e.g., Micrurus diastema) which
has resulted in a tangled taxonomic history
for several species.
The habitats of coral snakes also may be
useful for species identification, although
many species occur over several ecological
associations and may have elevational dis-
Dorsal aspect of Micrurus pachecogili (holotype, UTA R-12546). Total length = 639 mm.
tributions of over 2000 m. Nevertheless it is
worth noting that heretofore only a single
species of coral snake has been reported
from southern Puebla, namely M. Jaticollar-
is, a distinctive species and one of only two
species in Mexico possessing black body
rings arranged in triads. Micrurus laticollar-
is occurs on the west coast of Mexico in
Jalisco, Colima, and Michoacan and through
much of the Balsas Basin and associated
tributaries into southern Puebla where it may
be sympatric with M. pachecogili.
The distributions of a number of other
species of Micrurus, all with monadal black
body rings, approach the Zapotitlan Valley
within about 150 km, but are characterized
by markedly different habitats. The region
around Zapotitlan is dry, being in a rain-
shadow valley. This region receives about
250-350 mm of precipitation annually, and
the valley floor and surrounding slopes are
covered by arid scrub forest dominated by
many species of cactus. Micrurus bernadi
occurs in northern Puebla in tropical ever-
296
{a .
Ue
ea ee)
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Oaxaca
A Micrurus pachecoglili
@ Micrurus nebularis
Pacific Ocean
97°
Bie 33:
green forest and cloudforest at elevations of
50 to about 2000 m. There is some evidence
that this taxon is conspecific with M. dia-
stema (Pérez-Higareda & Smith 1990), also
known from the eastern versant of Mexico,
but this allocation has not been universally
accepted (Roze 1996). Micrurus browni is
known from the subhumid coastal tropical
forest, tropical deciduous forest, and dry
pine-oak forest of southern Mexico from
about sea level to over 2000 m. The ex-
tremely variable M. diastema inhabits trop-
ical evergreen forest and cloudforest of
eastern Puebla and adjacent Veracruz and
Oaxaca, as well as in subhumid forests over
96°
Distributions of two species of coral snakes, genus Micrurus, from the highlands of southern Mexico.
most of the Yucatan Peninsula. It ranges
from near sea level to about 1250 m. Mi-
crurus ephippifer zapotecus occurs in high-
land pine-oak forest of central Oaxaca at
elevations of 1700—2400 m; another appar-
ently closely related population, M. e.
ephippifer, occurs at lower elevations (100—
1500 m) in tropical deciduous forest and in
the ecotone of this forest with pine-oak for-
est. Micrurus nebularis occurs in pine-oak
forest at elevations of 2100—2300 m on the
southern slopes of the Sierra de Juarez in
central Oaxaca; all known specimens have
been collected in the vicinity of Ixtlan de
Juarez (Fig. 3).
VOLUME 113, NUMBER 1
During the course of my investigations
around Zapotitlan, I collected a series of
distinctive Lampropeltis triangulum which
subsequently became the type-series of a
new subspecies (Quinn 1983). This snake
closely resembles M. pachecogili, having
relatively long yellow body rings and nar-
row red rings, thus serving as one more
compelling example of mimicry.
Acknowledgments
I thank the Officials from the Direcci6n
General de Flora y Fauna Silvestre for is-
suing collecting permits for Mexico. How-
ard Arnott kindly took the photograph in
Fig. 2. For help at various times in the field,
I appreciate the efforts of Barry Armstrong,
William Lamar, and David Hillis.
297
Literature Cited
Campbell, J. A., & W. W. Lamar. 1989. The venomous
reptiles of Latin America. Cornell University
Press, Ithaca, 425 pp.
Pérez-Higareda, G., & H. M. Smith. 1990. The endem-
ic coral snakes of the Los Tuxtlas region, south-
ern Veracruz, Mexico.—Bulletin of the Mary-
land Herpetological Society 26:5—13.
Quinn, H. 1983. Two new subspecies of Lampropeltis
triangulum from Mexico.—Transactions of the
Kansas Academy of Science 86:113-—135.
Roze, J. A. 1996. Coral snakes of the Americas: bi-
ology, identification, and venoms. Krieger Pub-
lishing Company, Malabar, Florida, 328 pp.
Slowinski, J. B. 1995. A phylogenetic analysis of the
New World coral snakes (Elapidae: Leptomicru-
rus, Micruroides, and Micrurus) based on al-
lozymic and morphological characters.—Jour-
nal of Herpetology 29:325—338.
West, R. C. 1964. Surface configuration and associated
geology of Middle America. Pp. 33-83 in R.
Wauchope and R. C. West, eds., Handbook of
Middle American Indians, vol. 1, Natural En-
vironment and Early Cultures. University of
Texas Press, Austin, 570 pp.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):298-—301. 2000.
Fossil Red-shouldered Hawk in the Bahamas: Calohierax quadratus
Wetmore synonymized with Buteo lineatus (Gmelin)
Storrs L. Olson
Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.
Abstract.—A supposedly extinct genus and species of hawk, Calohierax
quadratus Wetmore, was originally described from a fragmentary tarsometa-
tarsus from Quaternary deposits in Little Exuma Island in the Bahamas. This
and a referred tibiotarsus from New Providence, Island, were later assigned to
the extant genus Buteo, but their specific identity remained uncertain. A pre-
viously unstudied humerus from a cave deposit on New Providence Island,
Bahamas, is here identified with the extant Red-shouldered Hawk, Buteo li-
neatus Gmelin, a species widespread in eastern North America and common
in peninsular Florida but unknown in the Bahamas. The other fossils are as-
signed to this species as well. Calohierax quadratus therefore becomes a syn-
onym of Buteo lineatus, which species has retreated from the Bahamas in the
late Quaternary for reasons that are unclear.
Among the severai new species of birds
that were described by Wetmore (1937)
from Quaternary cave deposits on Little
Exuma Island, Bahamas, was a new genus
and species of hawk, Calohierax quadratus,
based on the distal end of a tarsometatarsus.
The site was not on Great Exuma as Wet-
more originally reported (see Olson and
Pregill 1982:3). Brodkorb (1959) later re-
ferred the distal end of a tibiotarsus from a
cave deposit on New Providence Island,
Bahamas, to the same species. Restudy of
these specimens showed that the supposed
characters of the genus Calohierax were
founded entirely on artifacts of wear in the
case of the holotype, or intrageneric varia-
tion in the case of the referred specimen
(Olson & Hilgartner 1982). The specimens
were otherwise considered to be too frag-
mentary for specific identification, falling
within the range of size variation of the liv-
ing Red-shouldered Hawk Buteo lineatus
(Gmelin) or Gray Hawk B. nitidus (La-
tham).
Under present geographical and climatic
conditions, the Red-shouldered Hawk,
which occurs through most of eastern North
America and is abundant in Florida, would
be considered a more likely candidate for
having occurred in the Bahamas. The Neo-
tropical Gray Hawk now occurs only as far
north and east as Arizona and Texas. Nev-
ertheless, there is ample precedent for var-
ious Neotropical and western vertebrates
having occurred in eastern North America,
particularly Florida, in the Pleistocene (Em-
slie, 1998), so the possibility of Buteo ni-
tidus possibly having once occurred in the
Bahamas is not unthinkable.
The Bahaman fossil hawk was thus listed
simply as “‘“Buteo sp.’’, with the genus Ca-
lohierax Wetmore falling into the synony-
my of Buteo Lacepede, 1799 (Olson & Hil-
gartner 1982). The idea that there was an
extinct hawk in the Bahamas has lingered,
however, and Calohierax quadratus has
even appeared, in a reference that I can no
longer recall, under the absurd name
‘‘Quadrate Hawk.’’ Wetmore’s specific
name ‘“‘guadratus’’, it should be noted, was
derived from the squared appearance of one
of the tarsometatarsal trochleae, a feature
VOLUME 113, NUMBER 1
that turned out to be only an artifact of wear
(Olson & Hilgartner 1982).
Because of the ambiguities surrounding
the records of a medium-sized species of
Buteo in the Bahamas, it is fortunate that
an additional specimen bearing on its iden-
tity was located in the collections of the
Florida Museum of Natural History. This is
a nearly complete right humerus (UF
41801), lacking only a portion of the pec-
toral crest. It was collected by J. C. Dick-
inson and W. Auffenberg in the same “‘Ba-
nana Hole”? on New Providence Island that
yielded the fossils that formed the basis of
Brodkorb’s (1959) study and much of that
of Olson & Hilgartner (1982). It was col-
lected in 1958-1960, evidently after Brod-
korb’s study was completed, and since its
collection has apparently been overlooked.
The fossil humerus is in all details and
proportions identical with that in Buteo li-
neatus (Fig. 1) and in size falls squarely
among males from Florida (Fig. 2), which
belong to the southeastern subspecies B. /.
alleni Ridgway, which is smaller than the
nominate subspecies (Crocoll 1994). From
the small sample of Buteo nitidus exam-
ined, it is clear not only that B. nitidus is a
smaller species, with females in the range
of males of B. lineatus in length of the hu-
merus and the single male being much
smaller (Fig. 2), but the humerus is also
much more robust, the shaft especially be-
ing much thicker. Assuming that there was
only one species of Buteo in the Bahamas
in this size range, then the holotypical tar-
sometatarsus of Calohierax quadratus from
Little Exuma would belong to the same spe-
cies as the humerus from New Providence,
both islands being on the Great Bahama
Bank. Therefore the species Calohierax
quadratus Wetmore, 1937, becomes a ju-
nior subjective synonym of Falco lineatus
Gmelin, 1788.
The Red-shouldered Hawk is normally a
bird of moist riparian woods or swamp-
lands, hardly like the dry, scrubby habitats
that predominate in the Bahamas today.
Brown and Amadon (1968:578) remark that
299
this species “‘seems to be incompatible
with”’ the larger Red-tailed Hawk, B. ja-
maicensis (Gmelin), although this is as
much a reflection of the preference of the
latter for drier uplands rather than being due
to competition or antipathy. In any case, it
is the Red-tailed Hawk that occurs in the
Bahamas today, although it is an uncom-
mon resident only on some of the larger
northern islands and may perhaps be a re-
cent colonist, as it is absent in the fossil
record.
Prior to the arrival of Europeans only a
single terrestrial mammal lived in the Ba-
hamas, the hutia Geocapromys ingrahami
(Allen), which is now extinct on all but a
single small islet. The adults of this species
are too large to have served as prey for
Red-shouldered Hawks, but because these
hawks are very catholic in their choice of
food, taking birds, reptiles, amphibians,
large insects, and even crustaceans in ad-
dition to mammals (Crocoll 1994), and be-
cause all of these faunal elements are still
present in the Bahamas, it would be diffi-
cult to correlate the extinction of Buteo li-
neatus there to lack of suitable prey.
Habitat in most places in the Bahamas is
not now like that usually considered suit-
able for Red-shouldered Hawks, but is
thought to have been even more xeric in the
past (Pregill & Olson 1981, Olson & Hil-
gartner 1982). Thus, environmental and cli-
matic changes are also difficult to invoke as
an explanation for the disappearance of this
species from the Bahamas in the late Qua-
ternary.
The Bahaman fossil records constitute
the only evidence of Buteo lineatus any-
where in the West Indies, although B. ridg-
wayi of Hispaniola is now usually consid-
ered to be a derivative of B. lineatus (e.g.,
Sibley & Monroe 1990), so the Bahamas
may once have provided a stepping-stone
for this colonization.
Acknowledgments
I am most grateful to the staff of the Flor-
ida Museum of Natural History (UP),
300 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1. Right humeri of Buteo in anconal aspect: A, male Red-shouldered Hawk, B. lineatus, (UF 23893);
B, fossil referred to B. lineatus from New Providence Island, Bahamas (UF 41801); C, female Gray Hawk, B.
nitidus (UF 33746). Scale bar = 2 cm.
17
16
iis
14
humerus, distal width
1c
70 80 90 100
humerus length
Fig. 2. Scatter diagram showing length of humerus vs. distal width of humerus in male Buteo lineatus
(squares), female B. lineatus (triangles), the Bahaman fossil assigned to B. lineatus (star), female B. nitidus
(circles), and male B. nitidus (X). Five obviously missexed specimens have been corrected. All specimens of
B. lineatus are from Florida and are in the collections of the Florida Museum of Natural History.
VOLUME 113, NUMBER 1
Gainesville, for access to and loan of spec-
imens that made this study possible: David
W. Steadman and Tom Webber for modern
birds, and Marc Frank for the fossils. The
photograph is by John Steiner, Smithsonian
Photographic Services, and Fig. 2 was pre-
pared by Helen James.
Literature Cited
Brodkorb, P. 1959. Pleistocene birds from New Prov-
idence Island, Bahamas.—Bulletin of the Flor-
ida State Museum, Biological Sciences 4(11):
349-371.
Brown, L., & D. Amadon. 1968. Eagles, hawks, and
falcons of the world. 2 vols. McGraw Hill, New
York, 945 pp.
Crocoll, S. T. 1994. Red-shouldered Hawk Buteo li-
neatus.—Birds of North America 107:1—19.
Emslie, S. D. 1998. Avian community, climate, and
sea-level changes in the Plio-Pleistocene of the
301
Florida Peninsula.—Ornithological Mono-
graphs 50:1—113.
Olson, S. L., & W. B. Hilgartner. 1982. Fossil and
subfossil birds from the Bahamas. Pp. 22—56 in
S. L. Olson, ed. Fossil vertebrates from the Ba-
hamas.—Smithsonian Contributions to Paleo-
biology 48.
, & G. K. Pregill. 1982. Introduction to the pa-
leontology of Bahaman vertebrates. Pp. 1—7 in
S. L. Olson, ed., Fossil vertebrates from the Ba-
hamas.—Smithsonian Contributions to Paleo-
biology 48.
Pregill, G. K., & S. L. Olson. 1981. Zoogeography of
West Indian vertebrates in relation to Pleisto-
cene climatic cycles.—Annual Review of Ecol-
ogy and Systematics 12:75—98.
Sibley, C. G., & B. L. Monroe, Jr. 1990. Distribution
and taxonomy of birds of the world. Yale Uni-
versity Press, New Haven, 1111 pp.
Wetmore, A. 1937. Bird remains from cave deposits
on Great Exuma Island in the Bahamas.—Bul-
letin of the Museum of Comparative Zoology
80:427—441.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):302-—318. 2000.
Revision of the extant taxa of the genus Notiosorex
(Mammalia: Insectivora: Soricidae)
Leslie N. Carraway and Robert M. Timm
(LNC) Nash 104, Department of Fisheries and Wildlife, Oregon State University, Corvallis,
Oregon 97331-3803, U.S.A.; (RMT) Natural History Museum & Department of Ecology and
Evolutionary Biology, University of Kansas, Lawrence, Kansas 66045-2454, U.S.A.
Abstract.—We evaluated the taxonomic status of three specimens of gray
shrews, Notiosorex (Insectivora: Soricidae), collected in isolated mountain val-
leys in Tamaulipas, Mexico, with specimens referable to the Recent taxa N.
crawfordi crawfordi (n = 229) and N. c. evotis (n = 34). Statistical analyses
of cranial morphometric data revealed that the specimens from Tamaulipas
represented a heretofore unknown taxon which is described herein as a new
species of Notiosorex (N. villai). Also, N. c. evotis was found to be equally
distinct from N. c. crawfordi, thus was elevated to specific status.
Resumen.—Se evalu6 el estado taxonémico de tres especimenes de la mu-
sarana gris Notiosorex (Insectivora: Soricidae), colectados en valles dentro de
las motanas en Tamaulipas, México y referidos a los taxa recientes N. crawfordi
crawfordi (n = 229) y N. c. evotis (n = 34). Analisis estadisticos morfométricos
del craneo indicaron que los especimenes de Tamaulipas representan un taxon
desconocido, el cual se describe como una especie nueva de Notiosorex (N.
villai). Asimismo, N. c. evotis se encontr6é distinto a N. c. crawfordi, por lo
que se elevo al nivel especifico.
Gray shrews of the genus Notiosorex (In-
sectivora: Soricidae) are widely distributed
in the southwestern United States and
northern and western Mexico (Fig. 1). They
range from southern California, Nevada,
Utah, Colorado, Oklahoma, and western
Arkansas, southward to southern Baja Cal-
ifornia Sur, Michoacan, and southern Ta-
maulipas (Sealander 1952, Baker 1966,
Armstrong & Jones 1972, Woloszyn & Wo-
loszyn 1982, Carie et al. 1989). Insular re-
cords are known from Isla San Martin, Baja
California Norte (Schulz et al. 1970), and
Isla Palmito del Verde, Sinaloa, Mexico
(Armstrong & Jones 1971). Gray shrews
occur in a variety of habitats including de-
sert shrub (characterized by mesquites, Pro-
sopis; palo verde, Cercidium; Acacia; Yuc-
ca; Agave; and scattered Juniperus; Lange
1959), pine-oak forest (characterized by
Abies religiosa, Populus tremuloides, Jun-
iperus flaccida, four species of Pinus, and
three of Quercus; Alvarez 1963), in grass-
land with oak chaparral (characterized by
chamise, Adenostomna fasciculatum; scrub
oak, Quercus dumosa; California live oak,
Q. agrifolia; and mountain lilac, Ceano-
thus) and oak woodland habitats nearby
(Cunningham 1956), coastal sage scrub
(characterized by coastal sagebrush, Arte-
mesia californica; black sage, Salvia mel-
lifera; laurel sumac, Rhus laurina; and
grasses), yellow pine forest (Pinus austral-
is; Lange 1959), alkaline marsh (Stephens
1906), arid grasslands (containing scattered
catclaw, juniper, and mesquite; Baker
1966), and sandy flats (characterized by Ar-
temisia tridentata, Ephedra viridis, Peuce-
phyllum schottii, and Chrysothamnus vis-
cidiflorus; Fisher 1941). Elevations of oc-
cupied habitats range from 3 to 2618 m
(Fisher 1941, Lange 1959, Baker 1966, Da-
VOLUME 113, NUMBER 1
303
Fig. 1.
crawfordi (closed circles); 2, N. c. evotis (closed squares); and 3, Notiosorex unknowns from Tamaulipas (tri-
angles). Open circles, N. c. crawfordi, and open squares, N. c. evotis, represent collection localities for which
specimens were not examined (Bailey 1905; von Bloeker 1944; Cunningham 1956; Davis 1960; Baker 1966;
Dalby & Baker 1967; Armstrong 1972; Armstrong & Jones 1972; Findley et al. 1975; Sealander 1979; Hoff-
meister 1986; Caire et al. 1989; Rodriguez Vela 1999; Angeles Mendoza Duran, pers. comm.; Philip Myers,
pers. comm.).
vis & Sidner 1989). Because of the wide
range of habitats in which Notiosorex is
know to occur, the commonly used vernac-
ular name “‘desert shrew”’ (Hall 1981) is a
misnomer. Herein, we recommend the use
of gray shrew.
In 1953, Gerd H. Heinrich collected two
specimens of Notiosorex and in 1976
George D. Baumgardner collected a third
from western Tamaulipas. Findley (1955:
616) unequivocally categorized the former
two specimens as WNotiosorex crawfordi
crawfordi ‘‘on geographic grounds,”’
whereas Alvarez (1963:397) later referred
Distribution of gray shrews (Notiosorex) based upon specimens examined herein: 1, N. crawfordi
them to N. crawfordi noting that ““When
more abundant material is available the No-
tiosorex crawfordi of northeastern México
probably will be found to represent a new
subspecies.’” Schmidly & Hendricks (1984:
23) examined all three specimens and re-
ferred them to N. c. crawfordi because it
was “‘the name currently applied to desert
shrews in Texas and northern México.”’ As
these Tamaulipian shrews are isolated geo-
graphically from the widespread WN. c. craw-
fordi and are much larger and quantitatively
different from that taxon, we evaluated their
taxonomic status.
304
Materials and Methods
Specimens from throughout the distribu-
tion of Notiosorex (n = 266) were exam-
ined. Seven cranial and five mandibular
characters were recorded for each of the
139 specimens measured (Fig. 2). Relative
age of specimens was indexed by the lateral
length of the right I1 (first upper incisor;
Carraway et al. 1996). Quantitative char-
acters were analyzed by age to determine if
age caused a bias. Greatest length of skull,
rostral breadth, least interorbital breadth,
and cranial breadth were measured to 0.01
mm with Mitutoyo Digimatic electronic
calipers. All other quantitative characters
were measured by use of an ocular micro-
meter mounted in a Bausch and Lomb bin-
ocular microscope. Values were converted
from number of ocular lines to millimeters
for multivariate analyses and tabulation of
reported values. The states of a qualitative
character, roof of glenoid fossa extending
laterally from side of cranium, also were
recorded for each specimen.
Three a priori groups were formed of in-
dividuals of Notiosorex crawfordi crawfor-
di, N. c. evotis, and the three specimens
from Tamaulipas; assignment of individuals
to their a priori group was based on the geo-
graphic location of their collection site.
Multivariate analyses of the three a priori
groups were performed on the 12 quanti-
tative cranial and mandibular characters by
use of multigroup discriminant-function
analysis in BIOXTAT II (Pimentel 1995). A
large sample of N. c. crawfordi (n = 54)
from Huachuca Mts., Arizona, was exam-
ined for intraspecific geographic variation
and a comparison with the remaining N. c.
crawfordi was performed with multigroup
discriminant-function analysis. Univariate
and regression analyses were calculated by
use of STATGRAPHICS Plus (Statistical
Graphics Corporation 1995). For all analy-
ses, P < 0.05 was accepted as statistically
significant.
Standardized canonical vectors for the 12
quantitative variables were plotted on the
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
same graph as the three a priori groups with
the same canonical variate axes. The plot of
the variable vectors, when placed at the or-
igin of the canonical variates plot, provided
an indication of the direction and magni-
tude of effect that each variable had in dis-
criminating the three a priori groups (Joli-
coeur, 1959).
Summary statistics (CX SE. range, and
CV) were calculated for all 12 quantitative
variables for the three taxa of Notiosorex
under consideration (Table 1). Color of hair
in the pelage was determined by compari-
son with Munsell soil color charts (Munsell
Color 1975).
Results
Multigroup discriminant function analy-
sis of Notiosorex crawfordi crawfordi, N. c.
evotis, and Notiosorex unknowns from Ta-
maulipas, resulted in 99% correct classifi-
cation of individuals into their a priori
groups (Fig. 3). The two significant canon-
ical-variate axes (x7 = 153.61, df. = 24
and x”? = 37.73, df. = 11, respectively)
acounted for 81.10 and 18.90% of the var-
iation, respectively. Variance in the first
axis (canonical variate I) was accounted for
by greatest length of skull, 98.19%; cranial
breadth, 99.03%; length of unicuspid tooth-
row, 98.35%; width across M2-M2,
92.69%; length of mandible, 99.39%;
length of mandibular toothrow, 89.70%;
height of coronoid process, 96.08%; and
length of the coronoid process-ventral point
of upper condylar facet, 97.64%. The sec-
ond axis (canonical variate II) was affected
by length of the coronoid process-ventral
point of lower condylar facet (89.98%).
Only one individual (KU 105409), from 5
mi WNW EI Carrizo, Sinaloa, Mexico, was
not placed into its a priori group, WN. c. ev-
otis; it was classified as a N. c. crawford.
This individual also was classified as a N.
c. crawfordi based on morphological com-
parisons by Jones et al. (1962). Herein, this
individual is considered N. c. crawfordi for
purposes of further analyses.
VOLUME 113, NUMBER 1 305
glenoid
fossa
ar
4
11
9
10
LUC
Fig. 2. Camera-lucida tracing of skull of a Notiosorex (KU 145262) illustrating skull dimensions measured:
1, greatest length of skull; 2, rostral breadth; 3, least interorbital breadth; 4, cranial breadth; 5, length of maxillary
unicuspid toothrow; 6, length of maxillary complex toothrow; 7, width across M2—M2; 8, length of mandible;
9, length of mandibular toothrow; 10, height of coronoid process; 11, length of coronoid process-ventral point
of upper condylar facet; and 12, length of coronoid process-ventral point of lower condylar facet. Qualitative
character recorded is: 13, roof of glenoid fossa extending laterally from the side of the skull. Scale bar equals
5 mm.
306 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
CANONICAL VARIATE II
-6.2 -4.2
-2.2
1.8
-0.2
CANONICAL VARIATE |
Big. 3:
Canonical-variates plot of Notiosorex c. crawfordi (open circles), N. c. evotis (closed circles), and
Notiosorex new sp. (triangles) based on 12 measured variables. Canonical-variate axis I accounted for 81.10%
and canonical-variate axis II 18.90% of the variation present among the three taxa. Differences in cranial
morphology among the three taxa are characterized by the plot of the 12 variable vectors on the canonical-
variates plot in the upper right corner. Numbers are as in Fig. 2. The relationship of the variables among the
taxa are indicated by the length and orientation of the vector relative to the canonical-variate axes. From left to
right, taxa have shorter skulls relative to cranial breadth and taller and narrower coronoid processes. From bottom
to top, taxa have greater rostral breadth relative to length of maxillary complex toothrow.
Coefficients of variation for the N. c.
crawfordi from Huachuca Mts., Arizona,
ranged from 2.276% to 4.918% with one
outlier at 6.46%. Whereas, the same values
for the remaining N. c. crawfordi ranged
from 3.028% to 8.098%. This indicates
that among specimens of Notiosorex from
the Huachuca Mts. the level of variation
within and among the 12 quantitative char-
acters examined is very low. The one dis-
criminant function produced by a multi-
group discriminant-function analysis of the
two a priori groups of N. c. crawfordi was
not significant, at PP .< 0.05. Therefore:
these two a priori groups are acting as a
single unit.
Regression analyses indicated weak re-
lationships between age and length of uni-
cuspid toothrow for crawfordi (r? = 0.24)
and evotis (r? = 0.53) and between age and
least interorbital breadth (r? = 0.04) and
length of mandible (7? = 0.04) for crawfor-
di. When the discriminant analysis was re-
peated without these three variables, 93.5%
VOLUME 113, NUMBER 1
of individuals still were classified correctly
into their a priori groups.
For all N. c. crawfordi and N. c. evotis
the roof of the glenoid fossa extends later-
ally from the cranium. However, among the
Notiosorex from Tamaulipas the roof of the
glenoid fossa does not extend laterally from
the cranium.
Discussion
Morphometrically, the three specimens
of Notiosorex from Tamaulipas are most
similar to each other and are distinct from
all specimens of crawfordi and evotis (Fig.
3) in terms of size and shape of their skulls.
We had expected this outcome after making
careful visual comparisons (with aid of a
binocular microscope) of the three groups.
The discovery that evotis was equally dis-
tinct from the two other groups (Fig. 3) ne-
cessitated a reevaluation of its taxonomic
status. Previously, N. c. evotis was recog-
nized as being larger than N. c. crawfordi,
but we also found significant differences in
the shape of skulls and mandibles between
individuals of the two taxa.
Based on analyses reported herein we re-
fer the three specimens from Tamaulipas to
a new species and elevate N. c. evotis to
species level.
Class Mammalia Linnaeus, 1758
Order Insectivora Bowdich, 1921
Family Soricidae Fischer von Waldheim,
1817
Subfamily Soricinae Fischer von
Waldheim, 1817
Genus Notiosorex Coues, 1877
Notiosorex villai, new species
Villa’s Gray Shrew
Fig. 4A
Notiosorex crawfordi.—Findley 1955:616,
Ball & Kelson 1959:64 [part], Alvarez
1963:397, Hall 1981:65 [part], Schmidly
& Hendricks 1984:22.
Holotype.—Adult, female, skin and
skull; KU 54932, University of Kansas,
307
Natural History Museum, Mammal Collec-
tion; from “‘Jaumave, Tamaulipas, Mexico,
2400 ft.”’; obtained 26 July 1953 by Gerd
H. Heinrich, original number 7612. Alvarez
(1963:386) recorded the latitude and lon-
gitude of Jaumave as 23°34’N, 99°23’W.
Distribution.—Known only from the Po-
tosian Biotic Province (=Sierra Madre Ori-
ental Biotic Province) of the central moun-
tains of Tamaulipas, Mexico (Fig. 1).
Diagnosis.—As in all Notiosorex, speci-
mens of N. villai (Fig. 4A) have a combi-
nation of a deeply emarginated area be-
tween the condylar processes, e.g., interar-
ticular breadth about half the width of the
superior condylar process; the alveolus of
il extending posteriorly beneath at least
part of paraconid of ml (Carraway 1995);
pigment present of some teeth; and three
unicuspids.
Notiosorex villai can be distinguished
from other Notiosorex by the roof of the
glenoid fossa not extending laterally from
the cranium when the skull is viewed from
the dorsal aspect (Fig. 4A); and usually
from N. c. crawfordi by greatest length of
skull 216.97 mm and from N. c. evotis by
length of maxillary unicuspid toothrow
=2.0 mm, height of coronoid process <4.1
mm, length of coronoid process-ventral
point of upper condylar facet =3.8 mm, and
length of coronoid process-ventral point of
lower condylar facet =3.4 mm. All speci-
mens of N. villai can be separated from
specimens of N. c. crawfordi and N. c. ev-
otis by application of the following discrim-
inant-function equation: discriminant score
= 1.30722 (cranial breadth) — 0.34104
(height of coronoid process) — 0.01685
(ength of maxillary complex toothrow) —
4.87675 (least interorbital breadth) —
0.10742 (length of coronoid process-central
point of lower condylar facet) — 0.16575
(length of mandible) + 0.22460 (length of
mandibular toothrow) + 0.88996 (greatest
length of skull) + 0.22980 (length of max-
illary unicuspid toothrow) — 0.03089
(length of coronoid process-ventral point of
upper condylar facet) + 6.66117 (rostral
308
breadth) — 0.23311 (width across M2—M2)
— 11.0491. Those specimens with scores
=3.359 are referrable to villai and those
with scores =2.454 are referrable to craw-
fordi or evotis.
Description.—As in all soricids, Notio-
sorex villai has a double-faceted condylar
process and a fissident I1 with a large hook-
shaped anterior cusp and a posterior ven-
trally directed cusp. The skull is moderately
large, smooth, and without prominent ridg-
es and processes. The paroccipital processes
are small and lie against the exoccipitals
(Fig. 4A). The upper condylar facets are in-
flected; the corresponding areas in the su-
perior portion of the glenoid fossas are de-
pressed.
The pelage is composed of multibanded
hairs. In summer, hairs of the dorsal pelage
have a narrow band of silver gray (LOYR
6/1) distally and a wide band of dark gray
(7.5YR N4/O) proximally. Hairs of the ven-
ter are the same colors, except that the dis-
tal band of silver gray is wide and the prox-
imal band of dark gray is narrow producing
a silver wash. In winter, hairs of the dorsal
pelage have a narrow band of very dark
grayish-brown (1OYR 3/2) distally, a nar-
row band of pinkish white (7.5YR 8/2) me-
dially, and a wide band of very dark-gray
(7.5YR N3/O) proximally. Hairs of the ven-
tral pelage have a distal wide band of pink-
ish white (7.5YR 8/2) and a proximal nar-
row band of very dark-gray. The tail is very
dark grayish-brown.
Measurements.—Individuals of Notioso-
rex villai are smaller than those of N. c.
evotis, but larger than those of N. c. craw-
fordi (Table 1).
Ecology.—The three known specimens
of Notiosorex villai were collected in dif-
ferent habitats within the Potasian Biotic
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Province (Alvarez 1963). The area of Jau-
mave is pine-oak forest, that of Palmillas is
tropical forest, and the habitat of the col-
lection site near Rancho Carricitos is ripar-
ian. The subprovince, within the Potosian
Biotic Province, from which the three spec-
imens were collected contains ‘“‘low mese-
ta-like folded mountains” characterized by
“closely spaced ridges’”’ (Ferrusquia- Villa-
franca 1993:33).
Mammal associates of the holotype were
Baiomys taylori, Onychomys arenicola,
Oryzomys couesi, Peromyscus leucopus,
Reithrodontomys fulvescens, Sigmodon his-
pidus, Liomys irroratus, Mephitis ma-
croura, and Mustela frenata; only Pero-
myscus pectoralis was collected in the vi-
cinity of the specimen from Palmillas (G.
H. Heindrich, in litt.). Cryptotis parva,
Mormoops megalophylla, Desmodus rotun-
dus, Myotis californicus, Lasiurus borealis,
L. cinereus, Antrozous pallidus, Tadarida
brasiliensis, Sylvilagus floridanus, Sciurus
aureogaster, S. alleni, Liomys irroratus,
Peromyscus leucopus, P. pectoralis, P. boy-
lii, Baiomys taylori, Mus musculus, Bassar-
iscus astutus, and Mephitis mephitis were
collected in the vicinity of the specimen
from SW of Rancho Carricitos (Schmidly
& Hendricks 1984). One of the C. parva
was collected in the same pitfall as the No-
tiosorex. The greater species richness noted
for the latter locality is almost certainly the
result of greater trapping effort and not nec-
essarily an indication that the other mam-
mal associations were depauperate.
Etymology.—The species epithet is a pat-
ronymic to honor Bernardo Villa-R., the
‘‘father’”’ of Mexican mammalogy.
Remarks.—All three specimens of Notio-
sorex villai were collected in isolated
mountain valleys. The extent of the distri-
—
Fig. 4. Camera-lucida tracings of dorsal, lateral, and ventral views of the cranium, and lateral view of labial
side of mandible and oblique view of posterior portion of lingual side of left mandible of taxa of Notiosorex.
A, Holotype of Notiosorex villai (KU 54932, adult female). Note smooth lateral edge of cranium when viewed
from dorsal aspect (solid arrow), absence of extension of roof of glenoid fossa, small paroccipital processes
VOLUME 113, NUMBER 1 309
lying against the exoccipitals (open arrow), and coronoid process slender relative to height. B, N. evotis (KU
89214, adult male). Specimen from 17 km SW Choix, Sinaloa. Note prominent ridge on lateral edge of cranium
caused by extension of roof of glenoid fossa (solid arrow), low-set paroccipital processes extending at an oblique
angle from skull (open arrow), and coronoid process broad relative to height (dashed arrow). C, N. crawfordi
(KU 145262, adult male). Specimen from Peloncillo Mts., Guadalupe Canyon, Hidalgo Co., New Mexico. Note
prominent ridge on lateral edge of cranium caused by extension of roof of glenoid fossa (solid arrow), small
paroccipital processes lying against the exoccipitals (open arrow), and coronoid process slender relative to height
(dashed arrow). Scale bar equals 5 mm.
310
bution of the species has yet to be deter-
mined. However, considering the three dif-
ferent habitat types represented, N. villai
likely has a reasonably wide distribution in
Tamaulipas.
Heinrich, both in his field catalog and on
the specimen tag of one of the specimens
included herein, clearly recorded the local-
ity as Palmilla. However, on a map of Hein-
rich’s collecting localities drawn on 1 Sep-
tember 1953 by Hildegarde Heinrich, his
wife who accompanied him both in 1952
and 1953, the name is spelled Palmillas.
The hand-drawn map used by Heinrich in
1953 is believed to be based on the copy
of ‘“‘Mapa de la Republica Mexicana—Es-
tado de los Caminos Federales Estatales y
Vecinales’’ (1944) on deposit in the Natural
History Museum at the University of Kan-
sas. Notations made by E. R. Hall on the
latter map identify it as a gift to the muse-
um. The name of the town on this map is
spelled Palmillas; all subsequent authors
who have published upon this specimen
used the spelling Palmillas.
Specimens examined.—3, as follows.
Mexico: Tamaulipas: Jaumave, 2400 ft
(KU 54932, 2); Palmilla [sic], 4400 ft
[23°18’N, 99°33'W; Alvarez 1963:386]
(KU 54933, 2); 0.3 mi SW Rancho Carri-
citos, San Carlos Mts., 1900 ft (TCWC
30492, <).
Notiosorex evotis (Coues)
Large-eared Gray Shrew
Fig. 4B
Sorex (Notiosorex) evotis Coues, 1877:652.
N[otiosorex]. evotis Coues, 1877:652.
Notiosorex crawfordi evotis Merriam, 1895:
34; Hall & Kelson 1959:64 [part], Jones
et al. 1962:151, Hall 1981:65 [part].
Holotype.—Adult, no sex given, skin
only; USNM 9066, National Museum of
Natural History; from ‘“‘area of Mazatlan,
Sinaloa, Mexico’”’; obtained February 1868
by Ferinand Bischoff. The skull is men-
tioned by Coues (1877), but apparently was
never cataloged into the National Museum
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
collection. A sketch of the lateral view of
the right-side rostral area of the cranium
was included in Dobson (1890). However,
by the time Merriam (1895) published his
revision of the genus, the skull was missing.
Distribution.—Notiosorex evotis is
known from the states of Colima, Jalisco,
Michoacan, Nayarit, and Sinaloa, Mexico
(Fig. 1).
Diagnosis.—Notiosorex evotis can be
distinguished from N. villai by the roof of
the glenoid fossa extending laterally beyond
the curve of the cranium (Fig. 4B) and usu-
ally by length of maxillary unicuspid tooth-
row <2.1 mm, height of coronoid process
=4.3 mm, length of coronoid process-ven-
tral point of upper condylar facet =3.8 mm,
and length of coronoid process-ventral
point of lower condylar facet =4.0 mm; and
usually from WN. crawfordi by rostral
breadth =5.2 mm, width across M2—M2
=5.1 mm, length of mandible =7.4 mm,
height of coronoid process 24.3 mm,
length of coronoid process-ventral point of
upper condylar facet =4.0 mm, and length
of coronoid process-ventral point of lower
condylar facet =3.8 mm. N. evotis usually
can be distinguished from N. crawfordi by
use of a combination of height of coronoid
process and cranial breadth (Fig. 5). Based
on the results of the discriminant analysis,
all specimens of N. evotis can be separated
from specimens of N. crawfordi by appli-
cation of the following discriminant-func-
tion equation: discriminant score = 1.73654
(cranial breadth) + 0.22159 (height of cor-
onoid process) + 0.06177 (length of max-
illary complex toothrow) — 0.65656 (least
interorbital breadth) — 0.00471 (length of
coronoid process-central point of lower
condylar facet) + 0.27652 (length of man-
dible) + 0.08699 (length of mandibular
toothrow) — 0.86213 (greatest length of
skull) + 0.23251 (length of maxillary uni-
cuspid toothrow) — 0.02611 (length of cor-
onoid process-ventral point of upper con-
dylar facet) — 0.92248 (rostral breadth) +
0.10899 (width across M2—M2) — 37.2423.
Individuals with a score =—2.26 are refer-
VOLUME 113, NUMBER 1
rable to evotis and those with a score
=—1.65 are referrable to crawfordi.
Description.—Within extant members of
the genus Notiosorex, evotis has the largest
skull with prominent ridges (the roof of the
glenoid fossa) on the lateral sides of the cra-
nium and the paroccipital processes are set
low on the mastoid and extend at an oblique
angle from the skull (Fig. 4B). The coro-
noid processes are broad relative to their
height (Fig. 4B; Choate 1969:473, fig. 3b)
resulting in different placement of the con-
dyloid processes relative to each other
(Choate 1969:473, figs. 3b—c) for N. evotis
and N. crawfordi.
The pelage is composed of multibanded
hairs. In summer, hairs of the dorsal pelage
have a narrow band of very dark grayish-
brown (1OYR 3/2) distally, a narrow band
of pinkish white (7.5YR 8/2) medially, and
a wide band of dark gray (7.5YR N4/0)
proximally. The hairs of the venter have a
wide distal band of pinkish white and a nar-
row proximal band of very dark-gray
(7.5YR N3/0). The winter pelage consists
of hairs of the dorsum with a wide distal
band of very dark grayish-brown (1OYR 3/
2) and a proximal band of very dark-gray
(7.5YR N3/O). The hairs on the venter have
a wide distal band of light yellowish-brown
(1OYR 6/4) and a narrow proximal band of
dark gray (7.5YR N4/O). The hairs on the
tail are very dark grayish-brown.
Measurements.—Individuals of Notioso-
rex evotis are the largest members of extant
Notiosorex (Table 1).
Ecology.—Notiosorex evotis is known to
occur from 3-m elevation along the Pacific
Coast to 550 m in the Sierra Madre Occi-
dental, Sinaloa, and to 2317 m in the Sierra
Moroni, Zacatecas. It is know to occur in
habitats characterized by scattered cacti and
dense thornbush, and abandoned agricultur-
al fields bordered by an area of scattered
cacti, thornbush, and mesquite (scientific
names not given—Armstrong & Jones
1971); communities that “‘consisted mostly
of dry, dense weeds and short, thorny shrub
with a few trees” and “‘in low weeds near
311
thorn bush’’ (Jones et al. 1962:148—149);
‘‘*in damp spots under rocky ledges” (Fisher
& Bogan 1977:826); and in “‘semi-desert
habitat’? (Schlitter 1973:423).
Reported small-mammal associates are
Liomys pictus, Chaetodipus pernix, Sigmo-
don hispidus, and Mus musculus (Baker
1962, Jones et al. 1962).
Etymology.—The species epithet is de-
rived from the Greek ev, meaning good, and
otus, Meaning ear, possibly in reference to
its “‘extremely large’’ ears (Coues 1877:
652).
Remarks.—A discussion of the taxonom-
ic history of Notiosorex evotis is presented
in Jones et al. (1962).
Specimens examined.—34, as follows.
Mexico: Jalisco: 13 mi S, 15 mi W Gua-
dalajara (KU 33318); 21 mi SW Guadala-
jara (KU 42583-42585). Michoacan: 2 mi
E La Palma, SE side of Lago de Chapala
(KU 42586-42588). Nayarit: El Refilion
(USNM 508358); Tepic (USNM 314064).
Sinaloa: 20 km N, 5 km E Badiraguato (KU
96419); 16 km NNE Choix, 1700 ft (KU
89210-89213); 1 mi S El Cajoén, 1800 ft
(KU 100319); 15 mi SE Escuinapa (MSUM
5691); Isla Palmito del Verde, 6 mi NNW
Teacapan (KU 98880); Laguna, 17 km SW
Choix, 500 ft (KU 89214-89216); Maza-
tlan (KU 85533-85536; USNM 9066); 1 mi
N Mazatlan, 10 ft (MSUM 8149); 1 mi N
Mazatlan, 25 ft (MSUM 5690); Rosario,
500 ft (KU 90581); 10 km S, 38 km E Sin-
aloa (KU 125476—125479); 44 km ENE
Sinaloa, 600 ft (KU 89998); HWY 15, 0.25
mi S Sonora state line (UMMZ 109403).
Notiosorex crawfordi (Coues)
Crawford’s Gray Shrew
Fig. 4C
Sorex (Notiosorex) crawfordi Coues, 1877:
651.
Notiosorex crawfordi Coues, 1877:652,
True 1884:606, Merriam 1895:32, Hall &
Kelson 1959:64 [part], Hall 1981:65
[part].
Holotype.—Adult, no sex recorded, skin
312 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1.—Means (+SE), ranges (in parentheses), and CVs of measurements (in mm) of skull characters of
Notiosorex crawfordi crawfordi (n = 122), N. evotis (n = 14), and N. villai (n = 3).
Notiosorex
Character crawfordi evotis villai
Greatest length of skull 16.08 + 0.044 F206 O43 LF Le
(14.95—-17.25) (16.49-18.41) (16.97-17.27)<¢
0.03 0.03
Rostral breadth 4.83 + 0.02 3393" O04 5.19
(3.70-5.23) (5.0—5.63) (5.01—5.30)
0.04 0.03
Least interorbital 3.76 + 001 3.98 + 0.04¢ 3.83
breadth (3.42-4.14) (3.65-4.16) (3.57-4.01)
0.04 0.04
Cranial breadth 796. = 0.022 $567 O07? 8.30
(7.05—8.52) (8.02—8.99) (8.07—8.53)°
0.03 0.03
Length of maxillary 1:9 > 6:01 2.0 + 0.03 ae
unicuspid toothrow (1.5—2.2) (1.80—2.3) (2.0—2.3)
0.05 0.05
Length of maxillary 4.3 + 0.01 ABO =="0.03" 4.6
complex toothrow (3.9-4.6) (4.5—5.0) (4.44.7)
0.02 0.02
Width across M2—M2 4:3 + 0.01 DdyznO: 03° 5.0
(3.7-S.3) (4.9-5.4) (4.9-5.1)
0.02 0.02
Length of mandible 71.0°-= O02" 7.7 = 0.04 7.4
(6.4—7.7) (7.4-8.2) (7.3-7.5)°
0.03 0.03
Length of mandibular 4:7 = 0.014 5.0 2 0:03 5.1
toothrow (4.4-5.1) (4.9-5.3) (5.0-5.1)¢
0.02 0.02
Height of coronoid AOr= O:02 4720.05 4.1
process (3.44.9) (4.35.0) (4.0-4.1)°
0.02 0.04
Length of coronoid 3:3; HO102¢ 3.9. 0.04 3.8
process—ventral point (2.7-3.9) (3.84.2) (3.7-3.8)°
of upper condylar facet 0.06 0.03
Length of coronoid 347 70102 43° "O05 3.4
process—ventral point (3.2—-4.3) (4.0-4.7) (3.3-3.4)¢°
of lower condylar facet 0.05 0.05
«Sample size reduced by 6.
> Sample size reduced by 2.
©‘ Sample size reduced by 1.
4 Sample size reduced by 3.
and skull; USNM 2653/4437, National Mu-
seum of Natural History; from ‘‘near Fort
Bliss, about 2 miles above El Paso, El Paso
County, Tex.’’; obtained September 1861
by S. W. Crawford.
Distribution.—From southern California,
Nevada, Utah, Colorado, Oklahoma, and
western Arkansas, United States, southward
to southern Baja California Sur, and east-
ward to northern Sinaloa, and southern Za-
catecas and Nuevo Leén, Mexico (Fig. 1).
Diagnosis.—Notiosorex crawfordi can
be distinguished from WN. villai by the roof
of the glenoid fossa extending laterally be-
yond the curve of the cranium (Fig. 4C) and
by the greatest length of skull usually
<=16.98 mm. It usually can be distinguished
from N. evotis by rostral breadth <5.14
VOLUME 113, NUMBER 1
mm, width across M2—M2 <=5.1 mm, length
of mandible =7.5 mm, height of coronoid
process =4.4 mm, length of coronoid pro-
cess-ventral point of upper condylar facet
=3.9 mm, and length of coronoid process-
ventral point of lower condylar facet =4.1
mm. N. crawfordi usually can be distin-
guished from WN. evotis by use of a combi-
nation of height of coronoid process and
cranial breadth (Fig. 5). Specimens of N.
crawfordi can be separated from 100% of
specimens of N. villai and N. evotis by ap-
plication of the discriminant-function equa-
tions presented in the Diagnoses sections of
those species accounts.
Description.—The skull of Notiosorex
crawfordi is much like that of N. evotis, ex-
cept that it is much smaller, the coronoid
processes are slender compared to their
height, and the paroccipital processes are
small and lie against the exoccipitals (Fig.
4C). The latter two characters are similar in
form to N. villai. Although significant ge-
netic differences (M. B. O’Neill, C. Porter,
and R. J. Baker, pers. comm.) occur be-
tween populations of N. crawfordi in Baja
California and Texas, we found no identi-
fiable morphological differences.
The pelage is composed of multibanded
hairs. In summer, hairs of the dorsal pelage
have a narrow band of very dark grayish-
brown (1OYR 3/2) distally and a wide band
of dark gray (7.5YR N4/O) proximally. The
hairs of the venter have equal-width bands
of pinkish white (7.5YR 8/2) distally and
very dark-gray (7.5YR N3/0) proximally.
The winter pelage consists of hairs of the
dorsum with a narrow band of very dark
grayish-brown distally, a narrow band of
pinkish white medially, and a wide band of
dark gray (7.5YR N4/O) proximally. The
hairs of the venter have a wide band of
white (7.5YR N8/O) distally and a very nar-
row band of gray (7.5YR N5/0) proximally
creating a silvery wash effect over the ven-
ter. The hairs of the tail are dark grayish-
brown (1OYR 4/2).
Measurements.—Individuals of Notioso-
S13
rex crawfordi are the smallest extant mem-
bers of the genus (Table 1).
Ecology.—wNotiosorex crawfordi is
known to occur in habitats as diverse as de-
sert shrub and yellow pine forest (Lange
1959), pine-oak forest (Alvarez 1963), in
grassland with oak chaparral and oak wood-
land habitats nearby (Cunningham 1956),
alkaline marsh (Stephens 1906), sandy flats
(Fisher 1941, Yensen & Clark 1986), ‘‘arid
grasslands with scattered catclaw, juniper
and mesquite’? (Baker 1966:345), and near
‘“‘a mesquite tree on a moist mud flat”’
(Armstrong & Jones 1971:751). It is known
to occur at elevations at least as great as
2618 m (Davis & Sidner 1989).
From throughout the range of Crawford’s
gray shrew, known mammal associates are
members of the genera Cryptotis, Sorex,
Scapanus, Sylvilagus, Lepus, Tamias, Sciu-
rus, Spermophilus, Thomomys, Cratogeo-
mys, Liomys, Dipodomys, Perognathus,
Chaetodipus, Baiomys, Neotoma, Peromys-
cus, Onychomys, Reithrodontomys, Sigmo-
don, Microtus, Mus, Rattus, and Odoco-
ileus, plus 14 species of bats (Chiroptera;
Cunningham 1956, Anderson & Long
1961, Coulombe & Banta 1964, Baker
1966, Dalby & Baker 1967, Davis & Sidner
1989). Also, Crawford’s gray shrew is well
known for its association with woodrat
(Neotoma) nests throughout its distribution
(Armstrong & Jones 1972).
Etymology.—The species epithet is a pat-
ronymic to honor the collector of the type
specimen, S. W. Crawford.
Remarks.—Although trapping success
for specimens of Notiosorex crawfordi is
limited, remains thereof commonly occur in
pellets regurgitated by barn owls (Tyto
alba) and great horned owls (Bubo virgi-
nianus) throughout its distribution in the
United States and Mexico (Twente & Baker
1951, Baker 1953, Baker & Alcorn 1953,
Cunningham 1956, Anderson & Ogilvie
1957, Lange & Mikita 1959, Bradshaw &
Hayward 1960, Anderson & Long 1961,
Glass & Halloran 1961, Schaldach 1966,
Anderson 1972).
314
Jones et al. (1962:150—151), in a review
of Notiosorex from Sinaloa, treated evotis
and crawfordi as full species, stating that
‘“‘evotis has a longer body and hind foot
than crawfordi but a relatively (sometimes
actually) shorter tail and ear, and a distinct-
ly larger, heavier skull .... Notiosorex ev-
otis differs cranially from Notiosorex craw-
fordi as follows: larger . . . ; mesopterygoid
fossa squared rather than broadly U-shaped
anteriorly; rounded process on maxillary at
posterior border of infraorbital canal well
developed ...; occipital condyles smaller
and, in lateral view, elevated above basal
plane of skull; upper molars slightly more
crowded in occlusal view.”’ Armstrong &
Jones (1971:750), in their update on Sina-
loan mammals, treated the two forms as
subspecies stating that “‘the population in
northern Sinaloa is intermediate between
crawfordi and evotis’’ and that some of the
cranial differences noted earlier now appear
to be inconsistent. After examination of the
263 specimens referrable to N. crawfordi
and N. evotis included in this study, we
found that most of the characters presented
in Jones et al. (1962) are either inconsistent
or strongly age-related, thus are of limited
use in distinguishing crawfordi and evotis.
A discussion of the natural history, fossil
record, and reproduction of N. crawfordi
can be found in Armstrong & Jones (1972)
and Coulombe & Banta (1964).
Specimens examined.—229, as follows.
Mexico: Baja California: 9 mi N Catav-
ina on Mexico HWY 1 (MVZ 159725); 10
mi SE El Rosario (MVZ 159726); San Mar-
tin Island, 300 yds. inland from Hassler’s
Cave (MVZ 136207); San Quintin (CAS
52); San Quintin, San Simon River (USNM
139592-—139593); San Tomas (USNM
137142); Santa Anita (USNM 74550,
79088-79092, 146693, 146933-146934,
146936, 147352, 147421). Chihuahua: 3.5
mi ESE Los Lamentos (KU 76488); 2 mi
W Minaca (KU 109475). Coahuila: 3 mi
NW Cuatrociénegas (KU 51571-51572);
Sabinas (USNM 277621). Durango: 2 km
SE Atotonilco, 6680 ft (MSUM 13887-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
13888); 4.8 km SE Atotonilco, 6680 ft
(MSUM_ 13889-13890); 7 mi NNE Bo-
quilla, 6400 ft (MSUM_ 10260). Nuevo
Leon: 3 mi SW Galeana, 5100 ft (MSUM
11238). Sinaloa: On HWY 15, 0.25 mi S
Sonora line (UMMZ 109403); 5 mi WNW
El Carrizo (KU 105409); El Fuerte (KU
75184). Sonora: 14.6 mi E [by road] Ma-
zocahul (MVZ 148830); 4.1 mi NW [by
road] Nacori Chico (MVZ 148831). Zaca-
tecas: Plateado (USNM 90845).
United States: Arkansas: Crawford Co.:
Natural Dam (USNM 286549). Arizona:
Apache Co.: 1 mi N Spigerville (UMMZ
80236); Cochise Co.: 9.1 mi S Chiricahua
(USNM 552386); Huachuca Mts. (MSB
62141-62144, 62146-62148, 62150,
62154-62155, 62158-62159, 62161-—
62166, 62168;..62171-62173, 62173
62177, 62179, 62183-62184, 62188-
62189, 62210-62215, 62218-62219;
62223—62238);..20 mi E.. Pearce, -Pinery
Canyon, 6500 ft (UMMZ 64102); Coconino
Co.: 10 mi SW Black Falls (USNM
244129); Grand Canyon (USNM 250676);
Greenlee Co.: Blue River (USNM 144533);
Maricopa Co.: Phoenix South Mt. (CAS
13928); Pima Co.: 36 mi S Tucson (USNM
2725 15—272516); 40 mi S Tucson (USNM
272844); Santa Cruz Co.: locality unknown
(USNM 289955); Yuma Co.: Yuma
(USNM 120357). California: San Bernar-
dino Co.: San Bernardino (USNM 187011);
Inyo Co.: Panamint Range Cottonwood
Canyon (CAS 23228-23230); Saline Valley
(CAS 21270); Saline Valley, Grapevine
Canyon, 4036-5750 ft (CAS 23231-23232,
21249-21269); Silver Canyon Rd., 4.3 mi
E [by road] jct. with California HWY 6
(MVZ 158116); Riverside Co.: Millard
Canyon (CAS 23240); San Bernardino Co.:
Cottonwood (CAS 23242); Deep Canyon
(CAS 23238); Kingston Range (CAS
21518); San Gorgonia, Cottonwood (CAS
23233-23237, 23241); San Diego Co.: lo-
cality unknown (USNM 62619); Lakeside,
head of Wildcat Canyon, 2100 ft (KU
92627); Escondido (MVZ 33582); 9 mi S
Escondido (MVZ 33388); Santer Mts.
VOLUME 113, NUMBER 1
(USNM 62919). Colorado: Baca Co.: 14 mi
N, 4 mi E Springfield (KU 116960); Fre-
mont Co.: Phantom Canyon, Eightmile
Creek (KU 125348—125367); Wet Mt. (KU
125368—125379); Montezuma Co.: Mesa
Verde National Park (KU 105109); Otero
Co.: 3 mi NW Higbee (KU 51673). New
Mexico: Cibola Co.: Juan Tofoya (USNM
147966); Hidalgo Co.: locality unknown
(KU 145266, MSB 46468); 7.5 mi W An-
imas, Antelope Pass (KU 145258—145259);
Cienega Ranch ruins (KU 144031); 30 mi
E Douglas, Guadalupe Canyon, Peloncillo
Mts. (KU 145260—-145265); San Luis
Spring, Mexican Boundary (USNM
38250); Lincoln Co.: Capitan Mts. (USNM
127229); Shafer Ranch (UMMZ 114784);
Otero Co.: 3 mi N Tularosa (UMMZ
81380); Union Co.: Tollgate Canyon, 10 mi
N Folson (MWSU 15900). Nevada: Nye
Cor: 1 mi N, 5 mi E Grapevine, Peak Mt.,
5500 ft (MVZ 92391). Oklahoma: Cimar-
ron Co.: 2 mi N Kenton (MWSU 15867—
15868, 15895); 3 mi N Kenton (MWSU
15902); 4 mi N Kenton (MWSU 15779);
Comanche Co.: Wichita Mts. National
Wildlife Refuge (USNM 271959); Harmon
Co.: 4 mi S Hollis (OSU 5823). Texas: Ar-
cher Co.: intersection US 82. and 277, 22
mi SW Holliday (UMMZ 167208—167209);
14 mi WNW Archer City (MWSU 8584,
8586); Lake Kickapoo (MWSU 5614), NW
side Lake Kickapoo (MWSU 7016, 11106);
Bexar Co.: San Antonio (USNM 125708);
Brewster Co.: Burro Mesa, 3500 ft (MVZ
80281); Briscoe Co.: Tule Canyon (UMMZ
67277); Cottle Co.: 8 mi ESE Paducah (KU
64560); Dickens Co.: 1 mi E Dickens
(MWSU 2543); El Paso Co.: near Fort
Bliss, about 2 miles above El Paso (USNM
2653/4437); Garza Co.: locality unknown
(MMNH_ 12502-12503); 1 mi SE Post
(PSM 13878); Hansford Co.: 10 mi S, 3 mi
W Gruver (KU 119395); Howard Co.: Big
Spring (UMMZ 80248); Jim Wells Co.::
near Alice (TCWC 53283); Knox Co.: 4 mi
E Benjamin (MWSU 16023); Nueces Co.:
Corpus Christi (USNM_ 120087). Locality
unknown: (USNM 4437).
315
Conclusions
As presently understood, the geographic
distributions of these three taxa do not over-
lap, although those of Notiosorex crawfordi
and N. evotis are parapatric in northern Sin-
aloa. We believe the lack of overlap in geo-
graphic ranges is the result of these taxa
being low in abundance wherever they oc-
cur, use of trapping techniques inappropri-
ate for collecting shrews during surveys of
small mammals, and lack of collecting ef-
fort for soricids over large segments of the
distribution of Notiosorex in Mexico.
In multivariate space, not only can 100%
of N. villai, N. crawfordi, and N. evotis be
separated (Fig. 3), but N. villai can be dis-
tinguished from N. crawfordi and N. evotis
by the qualitative character of the roof of
the glenoid fossa not extending laterally
from the cranium (Fig. 4). Also, N. craw-
fordi and N. evotis can be distinguished by
the qualitative character of the relative
shape of the coronoid processes (Figs. 4B—
C; Choate 1969:473, figs. 3b—c) and the
quantitative relationship of height of coro-
noid process and cranial breadth (Fig. 5).
With recognition herein of three species
in the genus Notiosorex, the Mexican mam-
malian fauna now includes 28 species in the
family Soricidae (Ramirez-Pulido et al.
1996, Woodman & Timm 1999). In Mexi-
co, the greatest diversity of shrews is in the
Trans-Mexican Volcanic Belt and the Sierra
Madre del Sur provinces. Our discovery of
the distinctive new species Notiosorex villai
in Tamaulipas, part of the Sierra Madre Ori-
ental Province, an area not especially
known for endemism (Fa & Morales 1993),
and that all three species of Notiosorex oc-
cur in Mexico, highlights the need for con-
tinued collection of specimens and study of
available museum specimens to better un-
derstand the mammalian fauna of Mexico.
Acknowledgments
For the loan of specimens in their care,
we thank the curators and collection man-
agers of the California Academy of Scienc-
eS
oO
>
—~]
ae
iN
oie)
HEIGHT OF
>
CORONOID PROCESS (mm)
ee
Co
ee
on
7.0 7.4
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
O@o oO
o 6€6afa@O On
O GDOOKIEED ODOGEDD
@aoo ao
OQOOCCOGED @ O OO
00 0@m0
O qaimq@a0D00
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O O
O
7.8
@MO O
8.2 8.6 9.0
CRANIAL BREADTH (mm)
Fier:
Bivariate plot of height of coronoid process and cranial breadth illustrating almost complete separation
of Notiosorex evotis (closed circles) and N. crawfordi (open circles).
es (CAS); Michigan State University Mu-
seum (MSUM); Midwestern University,
Wichita Falls, Texas (MWSU); Mammal
Division, National Museum of Natural His-
tory (USNM); Collection of Vertebrates,
Oklahoma State University (OSU); Texas
Cooperative Wildlife Collection, Texas
A&M University (TCWC); Museum of
Vertebrate Zoology, University of Califor-
nia at Berkeley (MVZ); Museum of Zool-
ogy, University of Michigan (UMMZ);
James Ford Bell Museum of Natural His-
tory, University of Minnesota (MMNH);
the Museum of Southwestern Biology, Uni-
versity of New Mexico (MSB); and James
R. Slater Museum of Natural History, Uni-
versity of Puget Sound (PSM). We thank T.
Holmes for preparation of problematic
specimens, V. Sanchez-Cordero for trans-
lation of our Abstract into Spanish, and N.
A. Slade for statistical advice. EK Cervantes,
R. S. Hoffmann, V. Sanchez-Cordero, and
B. J. Verts read earlier drafts of this man-
uscript.
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):319-—333. 2000.
Status of the name Odocoileus hemionus crooki
(Mammalia: Cervidae)
James R. Heffelfinger
Arizona Game and Fish Department, 555 N. Greasewood Road, Tucson, Arizona 85745, U.S.A.
Abstract.—The present name of the desert mule deer (Odocoileus hemionus
crooki) is based on a specimen collected in southwestern New Mexico near
the Mexican border. This specimen was originally described as a new species
(Dorcelaphus crooki) of black-tailed deer, not as a mule deer, because many
of its characteristics are intermediate between mule deer and white-tailed deer.
In the same publication, Dorcelaphus hemionus eremicus is described from
western Sonora, Mexico, as a new subspecies of desert mule deer. A number
of mammalogists believed the type specimen of crooki to be a hybrid between
desert mule deer and Coues white-tailed deer (O. virginianus couesi), while
others hypothesized it represented extremes of normal variation in mule deer.
I have reassessed the type specimen of Dorcelaphus crooki and reaffirm that
it is a hybrid, invalidating the use of the crooki subspecies name. Consequently,
the oldest available name for the desert mule deer is O. h. eremicus.
Resumen.—E]1 nombre actual del venado bura del desierto (Odocoileus hem-
ionus crooki) se basa en un ejemplar colectado en el suroeste de Nuevo Mexico
cerca de la frontera con México. E. A. Mearns describi6 este eyjemplar como
una nueva especie (Dorcelaphus crooki) de venado cola negra, no como un
venado bura, debido a que muchos de sus caracteres son intermedios entre los
del venado cola blanca y los del venado bura. En la misma publicaci6n, Mearns
describio también a Dorcelaphus hemionus eremicus del occidente de Sonora,
México, como una nueva subespecie de venado bura del desierto. Un gran
numero de mastozodlogos ha considerado que el ejemplar tipo de crooki es un
hibrido entre el venado bura del desierto y el venado cola blanca de Coues (O.
virginianus Ccouesi), mientras que otros han hipotetizado que representaba ex-
tremos de la variaci6n normal del venado bura. Después de reevaluar el ejem-
plar tipo de Dorcelaphus crooki concluyo que es en realidad un hibrido. Por
lo tanto, el nombre valido mas antiguo para el venado bura del desierto es O.
h. eremicus.
Desert mule deer (presently known as
Odocoileus hemionus crooki Mearns, 1897)
inhabit the southwestern United States from
West Texas through southern New Mexico
and southern Arizona, and southward into
Sonora, Chihuahua, Coahuila, Zacatecas,
and Durango, Mexico (Leopold 1959, Cow-
an 1961, Wallmo 1981). In addition, a small
herd has been translocated to Nuevo Le6én,
Mexico (Morrison et al. 1992; Fig. 1).
Since its original description, the subspe-
cific name of this taxon has been conten-
tious due to uncertainties regarding the hy-
brid status of the type specimen.
Historical Review
Mearns (1897) described Dorcelaphus
crooki as a new species of black-tailed deer
because of its similarity to the Columbian
black-tailed deer (O. h. columbianus). The
type specimen (National Museum of Natu-
320
118°
0 100 300 kilometers
22°
118°
Big. 1.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
98°
ow
5
<=
8
at
fe)
Durango a
‘
* San Luis a
ee “¢ Potosi
Current distribution of desert mule deer in the southwestern United States and northern Mexico. Type
localities identified for a) Dorcelaphus hemionus eremicus Mearns, 1897; b) Dorcelphus crooki Mearns, 1897;
and c) Odocoileus hemionus canus Merriam, 1901.
ral History [USNM] 20572/35752) was col-
lected in 1892 by E. A. Mearns on the sum-
mit of the Dog Mountains, Hidalgo County
(formerly part of Grant Co.), New Mexico
during the survey of the boundary between
Mexico and the United States. No other
deer with similar characteristics were col-
lected at or near that locality.
In the same publication Mearns (1897)
described the ‘“‘Burro deer or desert mule
deer,’ Dorcelaphus hemionus eremicus
based on a male (USNM 63403, the type)
collected in 1895 ‘by. W. J..McGee in the
Sierra Seri, Sonora, Mexico. McGee did
not keep the skull, but processed the hide
as a deer skin rug. Mearns (1897) de-
scribed the subspecies as pale gray in color
with short pelage, a dark dorsal stripe, pal-
er forehead markings, and wide, heavy ant-
lers. This description was based on the
skin of the male and two sets of antlers,
each from different deer; one of which
came from the Sonoyta Valley, Sonora
(USNM 59910), the other from Black
Butte, Baja California (USNM 60855).
Pieces of skin trimmed from the hide as it
was made into a rug comprised the only
material representing the type specimen in
the National Museum until 1902 when the
rug was acquired from Anita McGee
(Poole & Schantz 1942).
In 1901, Merriam described Odocoileus
hemionus canus based on a male (USNM
99361) from Sierra en Medio, Chihuahua,
Mexico. Merriam (1901) distinguished O.
h. canus from the western subspecies O. h.
eremicus solely on the basis of antler con-
formation. The Sierra en Medio lies only 40
VOLUME 113, NUMBER 1
km southwest of the type locality for Dor-
celaphus crooki in New Mexico.
In his list of big game of North America,
and replacing Dorcelaphus with the correct
senior synonym QOdocoileus, Seton (1898:
286) included O. h. eremicus as a subspe-
cies of mule deer, but maintained the
‘Crook black-tailed deer”’ as O. crooki. Ly-
dekker (1915) also listed this animal as a
black-tailed deer, O. columbianus crooki.
Following Merriam’s (1901) description, O.
h. canus was used for the desert mule deer
in West Texas, Arizona, New Mexico, and
northcentral Mexico (Seton 1909, Lantz
1910, Lydekker 1915, Bailey 1931, Cowan
1936, Cahalane 1939, Dalquest 1953).
Several mammalogists believed that
crooki was based on a hybrid between
Coues white-tailed deer (O. virginianus
couesi) and a desert mule deer but lacked
known hybrids for comparison (Lydekker
1898, Seton 1929, Bailey 1931, O’Conner
1939). Goldman & Kellogg (1939) exam-
ined the holotype of O. h. canus and other
specimens from the Sierra en Medio, Chi-
huahua, along with the holotype of Dorce-
laphus crooki, and concluded that the type
of crooki was an unusual specimen of mule
deer and not a hybrid. Because the name
crooki antedates canus, they adopted O. h.
crooki as the correct name for desert mule
deer in the north-central states of Mexico
and adjacent Arizona, New Mexico, and
Texas. Hoffmeister (1962) also re-examined
the type of crooki and compared it with
specimens of O. virginianus and O. hem-
ionus. He interpreted the specimen as sim-
ply a mule deer with some features that
were intermediate with or shared by white-
tailed deer.
Based on the type specimen from Sierra
Seri, the range of O. h. eremicus was des-
ignated somewhat arbitrarily as western So-
nora, southwestern Arizona, and extreme
southeastern California (Mearns 1907:210).
Hoffmeister (1962) listed O. h. eremicus as
a synonym of O. h. crooki because he did
not consider western Sonoran mule deer
(burro deer) distinguishable from other pop-
321
ulations of desert mule deer. Only Cowan
(1936, 1961) and Longhurst & Chattin
(1941) attempted to quantify differences be-
tween deer within the ranges of eremicus
and crooki. Cowan’s (1936) interspecific
cranial distinctions were based on only four
eremicus Skulls; of the two additional er-
emicus specimens from Mexico that he
used to differentiate external characteristics,
one was from Tiburon Island, Sonora,
which Cowan (1961) later considered to be
a different subspecies (O. h. sheldoni).
Cowan (1961) based his differentiation of
eremicus On measurements of only one
male and one female specimen, which may
have been previously described (Cowan
1936, Longhurst & Chattin 1941). Long-
hurst & Chattin (1941) added descriptions
of pelage variations to differentiate eremi-
cus, but they had only one crooki skin and
three skulls for comparison. Cowan’s
(1936:236) measurements of eremicus from
southwestern Arizona and California are
within the normal variation of crooki re-
ported by Hoffmeister (1986). Hall (1981)
continued to treat western Sonoran mule
deer as a separate subspecies (O. h. eremi-
cus), but provided no supporting informa-
tion. Hoffmeister (1962) found mule deer
from southern Arizona and northern Sonora
within 80 km of the type locality of O. h.
eremicus (Sierra Seri, Sonora) to be suffi-
ciently similar to warrant treating eremicus
as a synonym of crooki. Hoffmeister
(1986), while remarking that he could not
confirm that O. h. eremicus was a synonym
of O. h. crooki, still implied that western
Sonoran mule deer were not distinguishable
from those farther east. The purpose of this
study is to confirm hybrid status of the type
specimen for O. h. crooki and clarify sub-
specific nomenclature for desert mule deer.
Material and Methods
I re-examine the type specimen of O. h.
crooki to compare and contrast its qualita-
tive and quantitative characters with corre-
sponding features of mule deer, white-tailed
322
deer, and their hybrids from southern Ari-
zona, southern New Mexico, and adjacent
Mexico. Published data from previous com-
parisons are supplemented by measure-
ments (in millimeters) from female O. h.
crooki (n = 12), female O. v. couesi (n =
17), the type specimen of Dorcelaphus
crooki, and a known O. h. crooki X O. v.
couesi F, hybrid. The type is an adult fe-
male, thus I included only adult females
(22 years old), as determined from tooth
wear and replacement (Robinette et al.
1957, Severinghaus 1949), for comparison.
Cranial measurements (Table 1) include the
six used by Hoffmeister (1962, 1986) in ad-
dition to depth of lacrimal fossa (deter-
mined as either shallow, deep, or “‘no de-
cision” by Hoffmeister 1962). All cranial
measurements were taken with a metric dial
caliper and recorded to the nearest 0.1 mm.
Values for paired measurements (e.g.,
length of right and left nasals) are averages.
Elsewhere in this report, I use the term hy-
brid to refer to only verified O. h. crooki X
O. v. couesi F, hybrids, unless otherwise
noted.
Review of Characters
Hoffmeister (1962) reviewed character-
istics useful for distinguishing O. virgini-
anus and O. hemionus in Arizona in his
evaluation of the type of crooki. These in-
cluded cranial and external measurements
(including size of metatarsal gland) along
with qualitative descriptions of the type of
antler, lacrimal pit, color pattern of the tail,
and color and position of the metatarsal
gland. He relied heavily on size because de-
sert mule deer are larger than Coues white-
tailed deer. Nevertheless, Hoffmeister did
not have adults of known hybrids available
to evaluate their size characteristics when
deciding on the taxonomic status of the type
of crooki. Although acknowledging Ni-
chol’s (1938) success in producing hybrids
in captivity, Hoffmeister (1962:52) tended
to discount the occurrence of hybrids in the
wild. Today, however, data on dimensions
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
and other characteristics are available for
mule deer X white-tailed deer hybrids, in-
cluding both captive produced and geneti-
cally confirmed, wild-taken animals. Char-
acteristics of known hybrids produced in
captivity provide a morphological basis for
assessing suspected hybrids between mule
and white-tailed deer encountered in the
wild. The primary features used to distin-
guish between desert mule deer and Coues
white-tailed deer concern the length and
color pattern of the tail; size, position, and
color of hair tuft of the metatarsal gland,
depth of the lacrimal pit, and dimensions of
the body and cranium (Table 1). My eval-
uation of the holotype of crooki based on
these features follows.
Metatarsal glands.—Metatarsal glands
of desert mule deer are positioned high on
the metatarsus, exceed 75 mm in length,
and are circumscribed with brown hair (Ca-
ton 1877, Hoffmeister 1986). Those of
Coues white-tailed deer are positioned be-
low the midpoint of the metatarsus, mea-
sure 25 mm or less in length, and are
rimmed by white hairs (Quay 1971, Hoff-
meister 1986). Unlike either parent, all
known F, hybrids have metatarsal glands
that are intermediate in length, location, and
appearance (Table 2). As affirmed by
Mearns (1907) and Bailey (1931), the po-
sition of the metatarsal gland in the holo-
type of crooki is intermediate in compari-
son to its location in white-tailed and mule
deer (Fig. 2). It is nearly identical in loca-
tion, length, and appearance to metatarsal
glands of F, hybrids produced in captivity
(Day 1980) and to wild mule X white-tailed
deer hybrids whose status was confirmed
genetically (Wishart 1980).
The metatarsal glands of the type of
crooki (right = 34 mm, left = 42 mm) are
longer than those of white-tailed deer (25
mm or less) and well below the range for
desert mule deer (75-150 mm; Table 1).
Day’s (1980) measurements of the metatar-
sal glands of two adult hybrids born in cap-
tivity were 50 mm for a female and 73 mm
for a male. Wishart (1980) reported lengths
VOLUME 113, NUMBER 1
of glands from two wild-taken mule X
white-tailed deer hybrids (status confirmed
by electrophoresis) as 50 mm for a female
and 62 mm for a male. Halloran & Kennedy
(1949) and Lang (1957:14) gave lengths of
metatarsal glands of adult female desert
mule deer from southern New Mexico that
averaged 2 to 3 times the length of the
gland in the holotype of crooki.
Anderson et al. (1964) included the co-
rona of hair (circumglandular hair tuft) in
their measurements of the metatarsal glands
of 431 adult female mule deer from the
Sacramento and Guadalupe mountains of
southern New Mexico. Metatarsal gland
lengths of females from the Sacramento
Mountains (identified as O. h. hemionus)
averaged 138 mm (range, 100—190); those
of the Guadalupe Mountains (identified as
O. h. crooki), 131 mm (range, 90-190). The
shortest (90 mm) is considerably longer
than the longest circumglandular hair-tuft
measurement (70 mm) on the type of
crooki.
Mearns (1897, 1907) described the hairs
surrounding the metatarsal gland of the type
of crooki as “‘sooty at the base and white
apically.”’ On examination, these hairs are
“sooty at the base;’’ however, while pale,
they are not white apically (Fig. 2c). Instead
they are nearly the same pale color as the
remainder of the leg, which Mearns (1897,
1907) described as ‘“‘cream-buff, except
where new clay colored hair is coming in
on the anterior border.’’ Metatarsal glands
of hybrids produced in captivity are either
circumscribed with white hairs (G. I. Day,
in litt.) or the hairs match the brown col-
oration of their mule deer parent (J. C.
Haigh, in litt.).
Tail.—Mearns (1897:2) described the tail
of crooki as “‘colored much as in D. col-
umbianus, but has a longer terminal switch;
upper side and extremity of tail all black,
lower side white medially, and naked to-
wards the base”’ (Fig. 3c). He gave its ver-
tebral length as 195 mm, which is longer
than that of a female hybrid (184 mm; G.
I. Day, in litt.) and in the range of O. v.
323
couesi, but at or exceeding the upper limit
for desert mule deer (Table 1). The color
pattern of the tail of the type of crooki re-
sembles the tails of captive-born hybrids,
which Day (1980) described as “‘dark red-
dish-brown or reddish-black above with
white beneath and along the borders”’ (Fig.
3d). Tails of some subspecies of mule deer
(e.g., O. h. fuliginatus) commonly have a
dark dorsal surface; however, this pattern is
rare in southern Arizonan and New Mexi-
can populations. Of 349 desert mule deer
observed in southeastern Arizona during
January 1998, no adults had dark tails re-
sembling the type of crooki. However, in
areas of sympatry with white-tailed deer,
two fawns seen in the company of female
mule deer each had a wide, dark tail stripe.
I do not know if these were hybrids or pure
mule deer fawns; yet hybrid fawns are typ-
ically seen in the company of mule deer
(Wishart 1980, Kay & Boe 1992), implying
that the usual hybrid cross is between an
aggressive white-tailed buck and a mule
deer doe.
Length of hind foot.—Hoffmeister (1962,
1986) allowed that the length of hind foot
(400 mm) of the crooki type is more char-
acteristic of white-tailed deer. Based on the
data at hand (Table 1), 400 mm is at the
upper extreme of length of hind foot for
Coues white-tailed deer and at the lower ex-
treme for desert mule deer. G. I. Day’s (in
litt.) measurement of the length of hind foot
in a captive-born hybrid is 405 mm.
Total length.—The total length of the
type of crooki (1440 mm; Mearns 1897)
and that of a female hybrid (1549 mm; G.
I. Day, in litt.) is within the range for desert
mule deer, but longer than normal for Coues
white-tailed deer (Table 1). The ratio of tail
length to total length for the type of crooki
is 7.4X which is at the upper extreme for
Coues white-tailed deer; however, the ratio
in Day’s female hybrid is 8.4 and within
the normal range for desert mule deer (Ta-
ble 1). The range of ratios of tail length to
total length in Coues white-tailed deer is
5.6X-—7.5X, whereas the normal range of
324 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1.—Comparison of Coues white-tailed deer (Odocoileus virginianus couesi), desert mule deer (O.
hemionus crooki), and an O. virginianus couesi X O. hemionus crooki F, hybrid, and the type specimen of
Dorcelaphus crooki. All measurements (in millimeters) are from adult females two years of age or older.
Characteristics of the type of Dorcelaphus crooki from Mearns (1907); cranial measurements from A. L. Gardner
(in litt.). Measurements of known hybrid (UA 22358) supplemented by morphological information from G. I.
Day (in litt.). Sample sizes in parentheses.
Interpretation
Character O. v. couesi F, hybrid Type of crooki O. h. crooki of crooki type
Metatarsal gland:
Location Below midpoint Intermediate Intermediate At or above mid- Hybrid
of shank? point of shank?
Length Usually <25? 50 34R, 42L™ 75-150? Hybrid
14—23 (4)? 102-121 (4)°
18 (1) X = 132°
Length hair tuft 70 90-190 (431)! Hybrid
Color hair tuft White Brown or white Pale brown Buff or Brown; Hybrid
never white*®
Tail:
Length 215-260 (3)8 184 195 170—228 (5) Mule deer or
Hybrid
>1888 145-180 (8)?
170-230 (9) 152-191 (5)
165-229 (8)!
127-185 (8)
Dorsal color White border, dull Dark reddish- Black White with black Hybrid
cinnamon? black? terminal brush"
Gray, reddish- Like white- No white border,
brown, grayish tailed deer‘ usually without
brown or almost midband?
black?
Length of hindfoot 387-390 (3) 405 400 430—464 (5)* Hybrid
<404° 380-490 (448)!
X = 409 406-445 (5)°
332—405 (18)4 406-475 (8)?
Total length 1410-1450 (3) 1549 1440 1370-1570 (5)* Mule deer or
1230-1420 (18) 1346-1549 (5)° Hybrid
1397-1702 (8)!
1430-1582 (8)!
Ratio of tail to total 5.6 X-6.6x (3)) 8.4 X TA X 6.0 X-8.9 x (5)* Hybrid
length 5.9 X-7.5 x (9) 8.2 <X-10.5 x™
6.4 X-7.3 x™ 8.1 X-10.2 x (5)<*
= ST Xe X = 8.0 X (8)
X = 10.1 x
Depth of lacrimal 3.0-5.7 (4)" 53 5.9Ro63L 6.4-11.2 (11)" Hybrid
fossa Shallow?" Deep*"
Length of ear Gilg pas 188 190 190-193 (2) Mule deer
145-170 (18) 184.2—203.2 (5)
175-220 (8)?
194.7—209.6 (8)!
Basilar length 205-216 (4)" DID 239-265 (7)" Hybrid
190.9-216.2 (13)° 229° 230-246 (12)
Length of nasals 71.3-79.8 (5)" 72.0 79.1—95.6 (8)*
60.4—76.7 (13)° 80.7° 79-95 (4)h Mule deer
56.4—61.0 (3) 62.5-81.7 (12)
Orbital width 55.3-59.7 (5)" 63 65.7—79.8 (10)" Hybrid
54.5—56.0 (4) 64.2° 68.0—81.0 (12)
VOLUME 113, NUMBER 1
Table 1.—Continued.
325
Interpretation
Character O. v. couesi F, hybrid Type of crooki O. h. crooki of crooki type
31.5=62.2 (13)°
51.4-60.8 (2)?
Zygomatic width 87.8—97.7 (6)* 96.3 99.1—113.7 (9)* Mule deer
90-97 (4) 102.1° 100-110 (13)
89.4—100.4 (11)°
96.6—100.7 (2)?
Length of upper P-M 63.5—67.2 (5)" 70.5 72.1-82.6 (11)? Mule deer
toothrow 61.1-69.6 (13)° 76.4° 80-89 (4)
63.4—-66.8 (4)* 70.1—85.5 (12)?
Length of lower P-M 72.5—76.2 (3)* 80.7 80.9—100.5 (10)" Mule deer
toothrow 66.1—77.6 (11)° S72" 87-97 (4)
72.1-—74.4 (2)? 82.4—97.4 (12)
Usual topographic 1231-21549 above 1800 <1400' White-tailed
elevation (m) deer
“Hoffmeister 1986.
> Quay 1971.
© Halloran & Kennedy 1949.
#<a. f. Day, in litt.
¢Lang 1959.
f Anderson et al. 1964.
® Caton 1877.
Mearns 1907.
‘Cowan 1961.
Pruett, im litt.
K Nichols 1938.
' Bailey 1931.
™ Hoffmeister 1962.
" This study.
° A. L. Gardner, in litt.
P Krausman et al. 1978.
4 Anthony & Smith 1977.
"Krausman 1978.
ratios in desert mule deer is 7.7X—10.5X
(Mearns 1907, Hoffmeister 1962, G. I. Day,
in litt.). An exception is a desert mule deer
from west of El Paso, Texas (Mearns 1907),
with an unusually long tail (228 mm) and
short total length (1307 mm) yielding a ra-
tio of 6.0X.
Length of ear.—Total length of ear for
the type specimen of O. h. crooki (190 mm)
is within the normal range for desert mule
deer and is longer than that of a white-tailed
deer (Table 1). The length of ear for two
adult F,; hybrids (O. h. crooki X O. v. coue-
si) was 188 mm (female), and 209 mm
(male), also within the normal range for
mule deer (G. I. Day, in litt.).
Lacrimal fossa.—Depth of the lacrimal
fossa is diagnostic; it is deeper (6.4—11.2
mm) and larger in desert mule deer than in
Coues white-tailed deer (3.0—5.7 mm).
Hoffmeister’s (1962:49) “tno decision’’ on
the depth of the fossa in the type specimen
of crooki was because both fossae are fe-
nestrate. However, the floor of each fossa is
clearly evident and the depth of the right
fossa measures 5.9 and the left fossa, 6.3.
These measurements are intermediate be-
tween the ranges of lacrimal-fossa depths of
the two species and confirms Wishart’s
(1980) observation for known mule X
white-tailed deer hybrids. Depth of lacrimal
fossa in another hybrid doe measures 5.3
(Table 1), which is near the upper range of
that for white-tailed deer.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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VOLUME 113, NUMBER | 327
Fig. 2. Metatarsal glands of: a) desert mule deer (Odocoileus hemionus crooki [=eremicus]); b) Coues white-
tailed deer (O. virginianus couesi); c) holotype of (Dorcelaphus crooki, USNM 20572/35752); d) known F,
hybrid (O. h. crooki X O. v. couesi). Photographs by J. R. Heffelfinger (a & b), L. M. Snyder (c), and G. I.
Day (d).
Fig. 3. Tails of: a) desert mule deer (Odocoileus hemionus crooki [=eremicus]); b) Coues white-tailed deer
(O. virginianus couesi); c) holotype of (Dorcelaphus crooki, USNM 20572/35752); d) known F, hybrid (O. h.
crooki X O. v. couesi). Photographs by J. R. Heffelfinger (a & b), L. M. Snyder (c), and G. I. Day (d).
328
Cranial measurements.—Hoffmeister’s
(1962, 1986) decision that the type of
crooki was a mule deer pivoted on the use
of skull measurements. Adult desert mule
deer in the region of the type locality of
crooki are much larger and may weigh
twice as much as Coues white-tailed deer
resulting in some cranial dimensions being
interspecifically diagnostic. Hoffmeister
(1962:48) provided generalized interspecif-
ic limits, but not actual ranges of the six
measurements (basilar length of Hensel,
length of nasals, orbital width, zygomatic
breadth, and length of both upper and lower
molariform toothrows) he used in separat-
ing female mule and white-tailed deer. He
said all six measurements of the type spec-
imen for crooki were within the lower range
of these measurements for desert mule deer.
However, I found that orbital width and
basilar length were below the correspond-
ing ranges for this measurement for mule
deer; the remaining four measurements
were within the normal range. The discrep-
ancy between Hoffmeister’s (1962) mea-
surement of the nasals of the type of crooki
and the longer measurement in Table 1
credited to A. L. Gardner results from Hoff-
meister’s measurement equaling the shortest
distance from the frontal-nasal suture to the
proximal margin of its anterior border be-
tween medial and lateral anterior projec-
tions. Gardner’s measurement is the average
of the greatest distance between anterior
and posterior points of right and left nasals;
the right nasal measures 80.0 mm and the
left, 81.5 mm. Hoffmeister’s (1986) princi-
pal components analysis using 11 cranial
measurements grouped measurements of
the type of crooki with those of mule deer.
Cranial measurements of an adult captive-
born hybrid doe are either intermediate or
within the normal range for mule deer (Ta-
ble 1).
Cowan (1962) reported four cranial di-
mensions (width of nasals, interorbital
width, palatal width, and postpalatal width)
used to differentiate mule deer and white-
tailed deer in Alberta, Canada. Three of
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
these four measurements from a male mule
x white-tailed deer hybrid (as determined
from metatarsal gland characteristics) were
within the normal range for mule deer. Wis-
hart (1980) used five cranial measurements
in an analysis of a male and a female wild-
taken hybrid (both confirmed by electro-
phoresis) in Alberta. All measurements fell
within the normal range for mule deer with
the exception of post-palatal width of the
female, which was within the range for
white-tailed deer.
Type locality.—Mearns (1897, 1907:190)
collected the type of Dorcelaphus crooki on
9 June 1892, on the summit of ‘Emory
Peak of the Dog Mountains, where I noted
its range as from 1500 to 1868 meters.”’
Mearns (1907:87) was camped at “Dog
Spring .. . [which] is about 2 kilometers...
north of [Boundary] Monument No. 55...
near the south extremity of the Dog Moun-
tains, a rugged range of which Emory Peak,
having an altitude of 1868 meters (6129
feet), is the highest.’”’ Dog Spring (31°21'N,
108°19'W) appears on several old maps as
“Ojo del Perro.’’ The Dog Mountains are
known today as the Alamo Hueco Moun-
tains. Apparently, the name Emory Peak
does not appear on any topographic map of
the area produced from 1881 through 1983
(C. Kollen, pers. comm.). A Department of
the Interior, General Land Office map dated
1903 shows an “Emory Sp.”’ northwest of
Ojo del Perro, in the general vicinity of
Pierce Peak (31°27'N, 108°20’W). As
Pierce Peak is drained to the north by Emo-
ry Canyon and 11 km northwest of Dog
Spring, it is most likely the same peak iden-
tified by Mearns as Emory Peak. The ele-
vation of Pierce Peak is given today as
1877 m; however, a map of the area dated
1942 has the elevation as 6149 ft (1874 m).
Mearns’ (1907) elevation of 1868 m for
Emory Peak is nearly equivalent; further-
more, Pierce Peak is covered with alligator
juniper (Juniperus deppeana), which
matches his description of the area.
The elevation at which the type of crooki
was collected, presumed to be above 1800
VOLUME 113, NUMBER 1
m, is above the normal upper elevational
limit for desert mule deer (1400; Krausman
1978, McCulloch 1972). Coues white-tailed
deer, however, often occur in highest den-
sities between 1230 and 2150 m elevation
(Anthony & Smith 1977). Both species are
present in the Alamo Hueco Mountains, but
white-tailed deer are less common today
than they were earlier in this century (Bai-
ley 1931; Raught 1967; A. Hurt, pers.
comm.).
Discussion
In the original description of Dorcela-
phus crooki, Mearns (1897:3) said ‘“The
skull has very nearly the same conforma-
tion as that of D. columbianus [black-tailed
deer], the lacrimal fossa being deeper than
in the Virginia deer, but shallower than in
the mule deer. The same intermediate con-
dition obtains with respect to the vomer, in
the relationships of the nasal and premax-
illary bones, in the form and size of the
teeth; and, in short, the whole animal ap-
pears to be a compromise between the char-
acteristics of the white-tailed and mule
deer’
In 1907, Mearns referred to this taxon as
Odocoileus crooki and explained naming
the deer for General George Crook. Mearns
also received a specimen of a 2-year-old
buck shot in the vicinity of Bill Williams
Mountain, Arizona, in 1884 by a member
of General Crook’s hunting party. Mearns
believed this specimen also represented his
new species of black-tailed deer. I have not
examined the male to verify its hybrid sta-
tus because its identity has no bearing on
the status of the name crooki because it is
not the type. Mearns (1907:187) reported
the length of metatarsal gland on this spec-
imen as 13 mm, which is typical of a white-
tailed deer (Table 1).
For several decades following Mearns’
(1897) description of Dorcelaphus crooki,
several authorities suggested that the type
was a hybrid (Lydekker 1898, Seton 1929,
Bailey 1931, O’Conner 1939) and Merri-
329
am’s name O. h. canus was used for the
desert mule deer. Goldman & Kellogg
(1939), having noted that a mule deer (O.
h. peninsulae) from lower Baja California,
Mexico, had a tail color pattern similar to
that of the type of crooki and reasoning that
their animal could not be a hybrid because
white-tailed deer were not in Baja Califor-
nia, concluded that the type of crooki was
an abnormal specimen of mule deer. Hoff-
meister (1962, 1986) acknowledged that the
type of crooki was intermediate in some
features, but believed the preponderance of
evidence, particularly of size, supported his
assessment that the animal was simply an
abnormal mule deer. He also said that the
metatarsal gland may be small or indistinct
in some populations of mule deer else-
where, citing Hershkovitz’s (1958:538) ob-
servation that two mule deer from lower
Baja California, Mexico, had poorly-devel-
oped glandular tissue underlying well-de-
fined metatarsal hair tufts. Nevertheless, the
type specimen of crooki has _ shortened
metatarsal glands and circumglandular tufts
unlike those of any known mule deer, but
consistent in size, form, and position with
those of known hybrids (Tables 1 & 2).
Hoffmeister (1962:52), in his statement
‘The few ‘hybrids’ that I have been able to
track down either prove to be clearly O.
hemionus or O. virginianus,’’ implied that
hybridization between these species proba-
bly did not occur in the wild.
White-tailed deer < mule deer hybrids
are known to have been produced in cap-
tivity as early as 1865 (Gray 1972). Other
examples of hybridization have been doc-
umented at captive facilities in Arizona (Ni-
chol 1938, Day 1980), Colorado (Spraker
et al. 1997), Illinois (Caton 1877), Texas
(Derr 1990), Wyoming (Guiroy et al. 1991,
E. S. Williams, in litt.), and Alberta, Canada
(Lingle 1992, W. D. Wishart, in litt.).
Whitehead (1972) reported white-tailed x
black-tailed deer hybrids produced in cap-
tivity in Tennessee.
Hybridization between white-tailed deer
and mule deer has been documented genet-
330
ically or on the basis of metatarsal gland
morphology in the wild in Arizona (Day
1964, P A. Dratch, in litt., J. A. Holcomb,
in litt.), Montana (Cronin 1991), Texas
(Carr et al. 1986, Stubblefield et al. 1986,
Derr 1990, Ballinger et al. 1992), Washing-
ton (Gavin & May 1988), Wyoming (Kay
& Boe 1992), and in Alberta (Wishart
1980) and British Columbia (Cowan 1962),
Canada. White-tailed deer and mule deer
are sympatric in the vicinity of the type lo-
cality of crooki (Hoffmeister 1962; A. Hurt,
pers. comm.), and hybrids are documented
from adjacent areas in Arizona and Texas.
The only genetic tests that will differen-
tiate white-tailed deer and mule deer are
electrophoresis of albumin (Scribner et al.
1984) and erythrocyte acid phosphatase (P.
A. Dratch, in litt.), and isoelectric focusing
of muscle esterase (Oates et al. 1979). All
of these analyses require fresh or frozen
samples. No molecular markers are cur-
rently known that will differentiate these
species from skin samples from museum
specimens (P. A. Dratch, in litt.).
Known hybrids are large and some body
(ength of ear and total length) and cranial
measurements (zygomatic breadth, length
of nasals, and upper and lower molariform
toothrows) are within the normal range for
desert mule deer (G. I. Day, in litt.). The
phenomenon of heterosis (hybrid vigor) in
F, hybrids is well known in cervids (Krzy-
winski 1993, Tate et al. 1997). The deer
farming industry has capitalized on heter-
osis by crossing the phenotypically diverse,
but presumed conspecific, red deer and wa-
piti (Cervus elaphus). F, hybrids show
higher and faster weight gains making them
more profitable than either purebred paren-
tal stock (Pearse 1993). Variability in over-
all size of F, phenotypes means that most
cranial and body measurements are poor
choices for evaluating hybrid status in deer
(Cowan 1962; Day 1980, in litt.; Wishart
1980).
Any mule deer may have an abnormal
tail, metatarsal gland, lacrimal fossa, cranial
measurements, or length of hind foot. How-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ever, many characteristics from the type
specimen for crooki fall outside the normal
range of variation for mule deer, and are
consistent with those of known white-tailed
x mule deer hybrids. The overwhelming
concordance of morphologic evidence in
this comprehensive analysis reveals the ho-
lotype of Dorcelaphus crooki as a hybrid
between Coues white-tailed deer and desert
mule deer. This, then, has serious repercus-
sions for the current scientific name for de-
sert mule deer. A scientific name based on
a type specimen later found to be a hybrid
is invalid and can not be used for either of
the parental species even if it has priority
over all other available names (ICZN 1985:
Art. 23[{h]). This reanalysis clarifies the
long-standing confusion regarding the sta-
tus of this contentious type specimen and
therefore the correct scientific name for this
taxon. The oldest available name for desert
mule deer (formerly known as O. h. crooki
and O. h. canus) is Odocoileus hemionus
eremicus Mearns, 1897; an abbreviated
synonymy follows:
Odocoileus hemionus eremicus (Mearns)
Dorcelaphus crooki Mearns, 1897:2; un-
available name because it is based on a
hybrid.
Dorcelaphus hemionus eremicus Mearns,
1897:4; type locality “Sierra Seri, near
the Gulf of California, in the most arid
portion of Sonora, Mexico.”
Odocoileus hemionus canus Merriam,
1901:560; type locality “‘Sierra en Media,
Chihuahua, Mexico.”’
Odocoileus hemionus crooki: Goldman &
Kellogg, 1939:507; name combination.
Acknowledgments
I am very much indebted to A. L. Gard-
ner, USGS Patuxent Wildlife Research Cen-
ter, Biological Survey Unit, National Mu-
seum of Natural History (formerly called
the United States National Museum), for
measurements of specimens including the
VOLUME 113, NUMBER 1
type of Dorcelaphus crooki and for tech-
nical advice and significant assistance with
the manuscript. L. M. Snyder of the Na-
tional Museum of Natural History provided
photographs of the type specimen of D.
crooki. Thanks are extended to V. C.
Bleich, D. E. Brown, M. A. Cronin, P. R.
Krausman, R. M. Lee, and J. R. Purdue for
their review of an earlier draft of this man-
uscript. EF J. Abarca, D. E. Brown, S. Gal-
lina, R. S. Henry, D. Humphreys, A. Mar-
tinez, E. Mellink, B. L. Morrison, J. Rom-
ero, A. Santos, E Tapia, M. Weber, and D.
L. Weybright provided important informa-
tion regarding the current range of desert
mule deer. Arizona Game and Fish Depart-
ment Wildlife Managers collected data on
tail color pattern during routine deer sur-
veys. J. C. deVos provided assistance with
photographs and M. Alderson generated the
map of the distribution of desert mule deer.
N. B. Carmony provided insightful com-
ments regarding subspeciation. I thank D.
E. Brown, P K. Devers, Y. Petryszyn, M.
T. Pruss, and especially R. S. White for al-
lowing access to specimens and for osteo-
logical assistance. G. I. Day, P. R. Kraus-
man... Y. McCulloch, and J. C. Truett
shared morphological data from their earlier
work. A. Fisher, A. Hurt, C. Kollen, C. R.
LeQuiev, E. M. Rominger, and C. G.
Schmitt provided details on the Dog/Alamo
Hueco mountains of southwestern New
Mexico. Thanks are also extended to E.
Mellink for providing the Spanish version
of the abstract. Production costs for this
manuscript were paid by the Federal Aid in
Wildlife Restoration Act, W-53-M and W-
78-R, of the Arizona Game and Fish De-
partment.
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Appendix |
Specimens Examined
With the exception of the two type specimens of
mule deer (both males) listed below, specimens used
in this analysis, including the type of Dorcelaphus
crooki, are all adult females (=2 years), and deposited
in the following institutions: Arizona State University
(ASU); Arizona Game and Fish Department, Tucson
(AGFD); University of Arizona (UA); National Mu-
seum of Natural History (USNM). A. L. Gardner mea-
sured the USNM specimens; I measured all others.
Odocoileus hemionus crooki [=eremicus| (14).—
United States. Arizona: Yavapai Co., 8 km NE Horse-
shoe Dam (ASU 643); Maricopa Co., 3 km E Horse-
shoe Dam (ASU 637); Pinal Co., N side of Canyon
Lake (ASU 643), Picacho Mountains (UA 24418-—
24420, 24429, 24430, 24436, 24478, 25299, 25308).
Mexico. Chihuahua: Sierra en Medio (USNM 99361,
type of O. h. canus). Sonora: Sierra Seri (USNM
63403, type of O. h. eremicus).
Odocoileus virginianus couesi (17).—United States.
Arizona: Cochise Co., Chiricahua Mountains (UA
20340, 20346); Graham Co., Blue River (USNM
32115); Pima Co., Santa Rita Mountains (AGFD 10,
UA 23304), Baboquivari Mountains (AGFD 53201);
Santa Cruz Co., Santa Rita Mountains (USNM
202931). New Mexico: Catron Co., Mogollon Moun-
tains (USNM 148574); Grant Co., 32 km W Silver
City (USNM 286685), head of Mimbres River (USNM
147476); Hidalgo Co., near Cloverdale (USNM
35748). Mexico. Chihuahua: Colonia Garcia (USNM
99347, 99350); Sonora: San Luis Mountains (USNM
36320); E side of San Luis Mountains (USNM 35751,
37085); Pozo de Luis (USNM 59229).
Odocoileus hemionus crooki [=eremicus] X O. vir-
ginianus couesi hybrids (2).—Arizona: Pima Co., Uni-
versity of Arizona Captive Facilities (UA 22358). New
Mexico: Hidalgo Co., Summit of the Dog Mountains
(USNM 20572/35752, type of Dorcelaphus crooki).
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(1):335-—336. 2000.
INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE
Applications published in the Bulletin of Zoological Nomenclature
The following Applications were published on 30 September 1999 in Vol. 56,
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Executive Secretary (I.C.Z.N.), % The Natural History Museum, Cromwell Road,
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Case No.
3116 Gnomulus Thorell, 1890 (Arachnida, Opiliones): proposed designation of G.
sumatranus Thorell, 1891 as the type species.
3089 Leucocytozoon (Protista, Haemosporida): proposed adoption of Berestneff,
1904 as the author and of Leukocytozoen danilewskyi Ziemann, 1898
as the type species.
Drosophila rufifrons Loew, 1873 and D. lebanonensis Wheeler, 1949 (cur-
rently Scaptodrosophila rufifrons and S. lebanonensis; Insecta, Dip-
tera): proposed conservation of the specific names by the designa-
tion of a neotype for D. rufifrons.
Vespertilio pipistrellus Schreber, 1774 and V. pygmaeus Leach, 1825 (cur-
rently Pipistrellus pipistrellus and P. pygmaeus; Mammalia, Chi-
roptera): proposed designation of neotypes.
Tanaecia coelebs Corbet, 1941 (Insecta, Lepidoptera): proposed conservation
of the specific name.
Diastylis Say, 1818 (Crustacea, Cumacea): proposed designation of Cuma
rathkii Krgyer, 1841 as the type species.
Hybognathus stramineus Cope, 1865 (currently Notropis strmineus; Ostei-
chthyes, Cypriniformes): proposed conservation of the specific
name
Strongylus tetracanthus Mehlis, 1831 (currently Cyathostomum tetracan-
thum) and C. catinatum Looss, 1900 (Nematoda): proposed con-
servation of usage by the designation of a neotype for C. fetra-
canthum
Musca geniculata De Geer, 1776 and Stomoxys cristata Fabricius, 1805 (cur-
rently Siphona geniculata and Siphona cristata; Insecta, Diptera):
proposed conservation of usage of the specific names by the re-
placement of the lectotype of M. geniculata by a neotype
Ichthyosaurus cornalianus Bassani, 1886 (currently Mixosaurus cornalianus;
Reptilia, Ichthyosauria): proposed designation of a neotype
Mystacina Gray, 1843, Chalinolobus Peters, 1866, M. tuberculata Gray, 1843
and Vespertilio tuberculatus J. R. Forster, 1844 (currently C. tub-
erculatus) (Mammalia, Chiroptera): proposed conservation of usage
of the names
336 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Holochilus Brandt, 1835, Proechimys J. A. Allen, 1899 and Trinomys Thom-
as, 1921 (Mammalia, Rodentia): proposed conservation by the des-
ignation of H. sciureus Wagner, 1842 as the type species of Holo-
chilus
Cervus gouazoubira Fischer, 1814 (currently Mazama gouazoubira; Mam-
malia, Artiodactyla): proposed conservation as the correct original
spelling
VOLUME 113, NUMBER 1 aor
Opinions published in the Bulletin of Zoological Nomenclature
The following Opinions were published on 30 September 1999 in Vol. 56, Part 3
of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be ob-
tained free of charge from the Executive Secretary, I.C.Z.N., % The Natural History
Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk).
Opinion No.
1930 Osilinus Philippi, 1847 and Austrocochlea Fischer, 1885 (Mollusca, Gastro-
poda): conserved by the designation of Trochus turbinatus Born,
1778 as the type species of Osilinus.
1931 Campeloma Refinesque, 1819 (Mollusca, Gastropoda): conserved.
1932 Holospira Martens, 1860 (Mollusca, Gastropoda): Cylindrella goldfussi Men-
ke, 1847 designated as the type species.
1933 Androctonus caucasicus Nordmann, 1840 (currently Mesobuthus caucasicus;
Arachnida, Scorpiones): specific name conserved.
1934 Paruroctonus Werner, 1934 (Arachnida, Scorpiones): conserved.
1935 Cicada clavicornis Fabricius, 1794 (currently Asiraca clavicornis; Insecta,
Homoptera): specific name conserved.
1936 Thamnotettix nigropictus Stal, 1870 (currently Nephotettix nigropictus; In-
secta, Homoptera): specific name conserved.
1937 Corisa propinqua Fieber, 1860 (currently Glaenocorisa propinqua; Insecta,
Heteroptera): specific name conserved.
1938 Musca rosae Fabricius, 1794 (currently Psila or Chamaepsila rosae; Insecta,
Diptera): specific name conserved.
1939 Trigonocephalus pulcher Peters, 1862 (currently Bothrops pulcher, Bothrie-
chis pulcher or Bothriopsis pulchra; Reptilia, Serpentes): defined by
the holotype, and not a neotype; Bothrops campbelli Freire Lascano,
1991: specific name placed on the Official List.
Hoplocephalus vestigiatus De Vis, 1884 (Reptilia, Serpentes): specific name
placed on the Official List.
Australopithecus afarensis Johanson, 1978 (Mammalia, Primates): specific
name conserved.
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Front cover—from this issue, p. 44.
CONTENTS
Austin Beatty Williams (17 October 1919—27 October 1999). Biographical summary
Rafael Lemaitre and Bruce B. Collette
A new species of mud shrimp, Upogebia cortesi, from Pacific Costa Rica (Decapoda: Thalassinidea:
Upogebiidae) Austin B. Williams and Rita Vargas
Periclimenes murcielagensis, a new species of shrimp (Crustacea: Decapoda: Palaemonidae) living on
black coral from the Pacific coast of Costa Rica Rita Vargas
A new squat lobster of the genus Munidopsis Whiteaves, 1874 (Crustacea: Decapoda: Galatheidae)
from Taiwan Ming-Feng Wu and Tin- Yam Chan
A new freshwater crab of the genus Geothelphusa Stimpson, 1858 (Crustacea: Decapoda: Brachyura:
Potamidae) from Yakushima Island, southern Kyushu, Japan Hiroshi Suzuki and Tomokazu Okano
A new species of the genus Neostylodactylus Hayashi & Miyake, 1968 (Crustacea: Decapoda:
Stylodactylidae) from southern Japan Junji Okuno and Hiroyuki Tachikawa
On the male of Scutumara enodis Ng & Nakasone, 1993 (Crustacea: Decapoda: Brachyura: Grapsidae)
N. K. Ng and T. Komai
Larval development of Cryptolithodes expansus Miers (Decapoda: Anomura: Lithodidae) reared in the
laboratory Mi Hyang Kim and Sung Yun Hong
A new genus of pinnotherid crab from the Indian Ocean (Crustacea: Decapoda: Brachyura)
Raymond B. Manning and Bella Galil
A new genus and species of ghost shrimp from Tobago, West Indies (Crustacea: Decapoda:
Callianassidae) Richard Heard and Raymond B. Manning
Griceus buskeyi, a new genus and species of calanoid copepod (Crustacea) from benthopelagic waters
off Hawaii Frank D. Ferrari and E. L. Markhaseva
Studies on the Crustacea of the Turks and Caicos Islands, British West Indies. IV. Heteromysis (Heteromysis)
spottei, a new species (Peracarida: Mysidacea: Mysidae) from Pine Cay
W. Wayne Price and Richard W. Heard
Gynodiastylis laciniacristatus, a new species (Crustacea: Cumacea) from Australia
Sarah Gerken and Jennifer Gross
Revision of the subterranean amphipod genus Spelaeogammarus (Bogidiellidae) from Brazil, including
descriptions of three new species and considerations of their phylogeny and biogeography
Stefan Koenemann and John R. Holsinger
Eudendrium bathyalis, a new species of hydroid (Hydrozoa: Anthomedusae: Eudendriidae) from
Bermuda Antonio C. Marques and Dale R. Calder
Cnidae of two species of Discosomatidae (Cnidaria: Anthozoa: Corallimorpharia) from Brazil
Suzana Machado Pinto and Maria Julia da Costa Belém
Additions to the cancellariid (Mollusca: Neogastropoda) fauna of South Africa
Richard E. Petit and M. G. Harasewych
Erpobdella lahontana (Annelida: Hirudinea: Arhynchobdellida: Erpobdellidae), a new species of fresh-
water leech from North America Peter Hovingh and Donald J. Klemm
A cladistic analysis of Sciomyzidae Fallén (Diptera) Luciane Marinoni and Wayne N. Mathis
Review of the chewing louse genus Abrocomophaga (Phthiraptera: Amblycera), with description of
two new species Roger D. Price and Robert M. Timm
A new species of the genus Cubacubana (Insecta: Zygentoma: Nicoletiidae) from a Mexican cave
Luis Espinasa
Three new species of bathyal cidaroids (Echinodermata: Echinoidea) from the Antarctic region
Rich Mooi, Bruno David, F. Julian Fell, and Thérése Choné
A new species of Pristigaster, with comments on the genus and redescription of P. cayana (Teleostei:
Clupeomorpha: Pristigasteridae) Naércio A. Menezes and Mario C. C. de Pinna
A new species of Apogon (Perciformes: Apogonidae) from the Saya de Malha Bank, Indian Ocean, with
redescriptions of Apogon regani Whitley, 1951, A. gardineri Regan, 1908, and A. heraldi
(Herre, 1943) Thomas H. Fraser
A new species of Pogonophryne (Pisces: Perciformes: Artedidraconidae) from East Antarctica
Richard R. Eakin and Arcady V. Balushkin
A new genus and species of inseminating fish (Teleostei: Characidae: Cheirodontinae: Compsurini) from
South America with uniquely derived caudal-fin dermal papillae
Luiz R. Malabarba and Stanley H. Weitzman
Calls and calling behavior of the frog Leptodactylus natalensis (Amphibia: Anura: Leptodactylidae)
W. Ronald Heyer and Celso Morato de Carvalho
A new species of venomous coral snake (Serpentes: Elapidae) from high desert in Puebla, Mexico
Jonathan A. Campbell
Fossil Red-shouldered Hawk in the Bahamas: Calohierax quadratus Wetmore synonymized with Buteo
lineatus (Gmelin) Storrs L. Olson
Revision of the extant taxa of the genus Notiosorex (Mammalia: Insectivora: Soricidae)
Leslie N. Carraway and Robert M. Timm
Status of the name Odocoileus hemionus crooki (Mammalia: Cervidae) James R. Heffelfinger
International Commission on Zoological Nomenclature
88
95
104
124
129
145
155
162
210
218
224
238
249
264
269
284
291
298
302
S19
335
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Taxonofriry~2
113(2):339-—355. 2000.
* ROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
fT evolutionary relationships of Phillips’ small-eared
-Shrew, Cryptotis phillipsii (Schaldach, 1966), from Oaxaca, Mexico
(Mammalia: Insectivora: Soricidae)
Neal Woodman and Robert M. Timm
(NW) Department of Biological Sciences, East Stroudsburg University, 200 Prospect Street,
East Stroudsburg, Pennsylvania 18301-2999, U.S.A.; (RMT) Natural History Museum and
Department of Ecology & Evolutionary Biology,
University of Kansas, Lawrence, Kansas 66045-2454, U.S.A.
Abstract.—The name Cryptotis peregrina (Merriam, 1895) previously en-
compassed two separate populations of a small-eared shrew of the Cryptotis
mexicana-group inhabiting the Sierra de Cuatro Venados and the Sierra de
Miahuatlan in Oaxaca, Mexico. Analysis of museum specimens from these two
populations, including the type series of C. peregrina, indicates that they are
neither conspecific nor even sister taxa. We taxonomically restrict the name C.
peregrina to populations in the Sierra de Cuatro Venados and adjoining Sierra
Yucuyacua, whereas the name Cryptotis phillipsii (Schaldach, 1966) is avail-
able for the population in the Sierra de Miahuatlan. Moreover, we demonstrate
that C. phillipsii is the most primitive member of the C. mexicana-group,
whereas, C. peregrina is one of the more derived members of the group.
Choate’s (1970) comprehensive revision
of the small-eared shrews of the genus
Cryptotis divided the Mexican and Central
American species into three informal
groupings: the ‘‘Cryptotis mexicana-
group,” the “‘Cryptotis parva-group,”’ and
a group of three “relict species.”’ At that
time, C. mexicana (with four subspecies),
Cryptotis goldmani (with two subspecies),
and the monotypic Cryptotis goodwini
comprised the C. mexicana-group. Based
on cranial and postcranial morphology (in
particular, unique modifications of the fore-
limb) and biogeographic patterns, we
(Woodman & Timm 1999) recently revised
the C. mexicana-group, providing evidence
that it may represent a natural grouping dis-
tinct from other members of the genus. Our
revision recognized eight species in the C.
mexicana-group, many of which were treat-
ed previously as subspecies (Choate 1970,
Hall 1981): C. alticola, C. goldmani, C.
goodwini, C. griseoventris, C. mexicana, C.
nelsoni, C. obscura, and C. peregrina.
Cryptotis peregrina, which is endemic to
Oaxaca, Mexico, was first described by
Merriam (1895) as a subspecies of C. mex-
icana based on a series of 24 specimens
collected by E. W. Nelson and E. A. Gold-
man in 1894 ‘from mountains 15 miles
[south] west of city of Oaxaca”’ (Merriam
1895:24). Binford (1989) identified this
range of mountains as the Sierra de Cuatro
Venados. In addition to the type series,
Choate (1970) referred specimens from the
Sierra de Miahuatlan in southern Oaxaca,
including the holotype and two paratypes of
Notiosorex phillipsii, to C. mexicana pere-
grina. We initially agreed with Choate’s as-
sessment, referring material from the Sierra
de Cuatro Venados and the Sierra de Mia-
huatlan to the species Cryptotis peregrina
(Woodman & Timm 1999). Cranially, the
shrews previously considered by Choate
(1970) and subsequent authors (e.g., Hall
340
1981) to be subspecies of C. mexicana (C.
mexicana, C. nelsoni, C. obscura, C. pere-
grina) are quite similar and can be difficult
to distinguish. However, the foreclaws vary
noticably in length and breadth among
members of the C. mexicana-group (Choate
1970, Woodman & Timm 1999), and we
showed that other aspects of the forelimb,
in particular the morphology of the humer-
us, are distinct as well (Woodman & Timm
1999). Our phylogenetic analysis indicated
that C. peregrina was the most plesio-
morphic member of the C. mexicana-group.
Our study (Woodman & Timm 1999) fo-
cused mainly on the larger, broad-clawed
shrews that we termed the Cryptotis gold-
mani-group, which are a subset of the C.
mexicana-group. Initially, we were interest-
ed in the former subspecies of C. mexicana
primarily as outgroups to aid in polarizing
characters for phylogenetic analysis. In
studying Cryptotis peregrina, we depended
heavily on specimens from the Sierra de
Miahuatlan in the University of Kansas
Natural History Museum collection, be-
cause they were readily available to us and
included the only postcranial material then
known for the species. One inconsistency
we noted between our study and Choate’s
revision was that he referred to C. [m.] mex-
icana as “having less highly developed
front feet and claws’? (Choate 1970:232)
than C. [m.] peregrina. However, the spec-
imens from the Sierra de Miahuatlan had
distinctly shorter and narrower claws than
C. mexicana. We also observed that their
humeri were less modified that those of C.
mexicana, adding to our view of their being
more plesiomorphic.
Recently, we re-studied the holotype and
type series of Cryptotis peregrina at the Na-
tional Museum of Natural History and dis-
covered that Choate’s (1970) observations
were correct—specimens in the type series
from Sierra de Cuatro Venados all possess
longer, broader claws than C. mexicana. In
addition, we were graciously granted per-
mission by the American Museum of Nat-
ural History to remove the humerus from
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
the only known fluid-preserved specimen of
C. peregrina from the Sierra Yucuyacua, a
mountain range in western Oaxaca that ad-
joins the Sierra de Cuatro Venados. Study
of these specimens indicates that the two
populations of Cryptotis inhabiting the Si-
erra de Cuatro Venados\Sierra Yucuyacua
complex and the Sierra de Miahuatlan, re-
spectively, are not conspecific. Herein, we
taxonomically restrict the name Cryptotis
peregrina to shrews inhabiting the former
region. The name Cryptotis phillipsii
(Schaldach, 1966) is available for the pop-
ulation in the Sierra de Miahuatlan. Much
of what we reported earlier in regard to C.
peregrina (Woodman & Timm 1999) actu-
ally refers to a combination of C. peregrina
and C. phillipsii. It is our purpose here to
redescribe C. peregrina and C. phillipsii in
light of this new information and to clarify
the distinctions among the species in the
Cryptotis mexicana-group. In addition, we
present a refined hypothesis of phylogenetic
relationships among the species in this
group of shrews.
Methods
Our current work used the techniques
and methods of our previous studies of
shrews (Woodman & Timm 1992, 1993,
1999), and more detailed accounts of meth-
odology are presented there. Species and
subspecies synonomies list only published
uses of names. Regional names, place
names, and coordinates in Oaxaca derive
from Binford (1989), Choate (1970), and
our review of maps and gazetteers of the
region. Terminology of dentition and dental
characteristics follows Choate (1970). An-
atomical terminology of the humerus and
other aspects of the postcranial skeleton fol-
lows Reed (1951). Measurements used in
our analyses follow Woodman & Timm
(1993, 1999); abbreviations of measure-
ments are explained in Table 1. Measure-
ments of the skull were taken to the nearest
0.1 mm using either an ocular micrometer
in a binocular microscope or a hand-held
VOLUME 113, NUMBER 2
dial caliper (for condylobasal-length and
cranial-breadth). Univariate statistics in-
clude mean + standard deviation.
Multivariate analysis was carried out us-
ing Minitab 8.0. Our principle components
analysis (PCA; Fig. 1, Table 2) used ten
logarithm-transformed cranial measure-
ments (condylobasal-length, breadth-of-zy-
gomatic-plate, interorbital-breadth, breadth-
across-U's, breadth-across-U?s, breadth-
across-M?’s, palatal-length, length-of-upper-
toothrow, length-of-unicuspid-toothrow,
length-of-upper-molariform-toothrow) from
12 Cryptotis peregrina and 14 C. phillipsii
with complete crania.
Phylogeny and character evolution were
analyzed using PAUP 4.0b2 (see Swofford
1998) and MCCLADE 3.0 (see Maddison
& Maddison 1992). Phylogenetic analysis
(Fig. 2) was carried out using an exhaustive
search of 32 unordered and equally-weight-
ed transition series (Table 3, Appendix I),
with Cryptotis parva parva and C. nigres-
cens serving as outgroups for polarizing
characters.
Specimens from the following institu-
tions were used in this study: American
Museum of Natural History, New York
(AMNH); Natural History Museum, Lon-
don (BM); California Academy of Scienc-
es, San Francisco (CAS); Escuela Nacional
de Ciencias Biolédgicas, Mexico City
(ENCB); Instituto de Biologia, Universidad
Nacional Aut6noma de México, Mexico
City (UNAM); University of Kansas Natu-
ral History Museum, Lawrence (KU); Nat-
ural History Museum of Los Angeles
County, Los Angeles (LACM); Museum of
Comparative Zoology, Cambridge (MCZ);
James Ford Bell Museum of Natural His-
tory, St. Paul (MMNH); Museo de Zoolo-
gia, Facultad de Ciencias, Universidad Na-
cional Autonoma de México, Mexico City
(MZFC); Texas Cooperative Wildlife Col-
lection, College Station (TCWC); Univer-
sity of Michigan Museum of Zoology, Ann
Arbor (UMMZ); National Museum of Nat-
ural History, Washington (USNM).
341
Results of Multivariate Analysis
Principle components analysis (PCA) of
cranial variables supports the separation of
Cryptotis peregrina and Cryptotis phillipsii.
A plot of factor scores on factor axes | and
2 shows a clear distinction between these
two species with minimal overlap (Fig. 1).
This plot emphasizes the generally larger
cranial size (factor axis 1—Table 2) of C.
phillipsii, despite the generally similar ex-
ternal measurements of the two species (Ta-
ble 1). In the region of size overlap between
the two species on the plot, C. peregrina
and C. phillipsii are separated by the com-
bined effects of the two axes, suggesting a
common, but offset trend of decreasing uni-
cuspid-toothrow-length and palatal-length
and increasing interorbital-breadth and pal-
atal-breadth (M2B) with increasing size
(Table 2).
Revised Phylogeny of the
Cryptotis mexicana-group
Our re-analysis of phylogenetic relation-
ships within the Cryptotis mexicana-group,
based on 32 transition series (Table 3), re-
sulted in a single most parsimonious tree of
67 steps (Fig. 2). Discounting differences
such as the exclusion of Cryptotis goodwini
magnimana and the inclusion of Cryptotis
phillipsii as a species separate from Cryp-
totis peregrina, the new tree essentially
matches three of the nine most parsimoni-
ous trees from our previous phylogenetic
analysis of this group of shrews (Woodman
& Timm 1999, fig. 13, central column of
trees). The four species that comprised the
Cryptotis goldmani-group (C. alticola, C.
goldmani, C. goodwini, C. griseoventris)
form the crown clade of the C. mexicana-
group, whereas the species previously sub-
sumed under the name C. mexicana (C.
mexicana, C. nelsoni, C. obscura, C. pere-
grina, C. phillipsii) appear as individual
branches at different levels of the tree. This
supports our previous view that C. mexi-
cana (sensu latu) was paraphyletic with re-
spect to the C. goldmani-group.
342 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1.—Measurements of Cryptotis mexicana-group shrews used in this study. Statistics presented are mean
+ standard deviation of the mean, and observed extremes.
C. mexicana
(Oaxaca) C. nelsoni C. obscura C. peregrina C. phillipsii
Skin measurements
n= 118 n=9 n= 55 n= 24 n=27
Head and body length (HB)
69 + 4 16.23 66 = 4 F2-s03 68 + 6
55-88 72-79 56-75 66-78 58-78
Tail length (TL)
Of Gracies DX NaS 26s o 30) E72 Sas ¥3
20-34 22-33 20-32 24-33 23-36
Craniomandibular measurements
n = 30 n=6 n = 32 n= 12 n= 14
Condylobasal-length (CBL)
18.7 + 0.4 L9:5.250)3 18.4 + 0.4 19.1 +-04 1952 05
17.5-19.3 19.1—20.0 17.4-19.1 18.3-19.8 18.8—20.2
Breadth-of-braincase (BB)
9:82 0:3 104 43.033 916. > 02 9:0°> 0.1 10.3 =O
9.3-10.7 9.9-10.7 9.2—10.2 9.7-10.1 9.8-10.7
(n = 28) (n = 30) (n = 11) (n = 13)
Breadth-of-zygomatic-plate (ZP)
LS oy 220622 (Oe 2.0) 25 Ost 1.6-2,04 20.70%
1.42.0 1.0—2.2 1.8—2.1 1.5—-1.8 1.8—2.3
Interorbital-breadth(IO)
A.J 102 5. OH ry 0.1 A Tea b 419) =O
4.3-5.0 4.8-5.2 4.44.9 4.64.9 4.75.1
Breadth-across-first-unicuspids (U1B)
DAO 28 Of 25041 yes vagal 0) | 26° = Ol
2.3—2.5 2.6—2.9 2.3—2.6 2.1—2.6 2.5—2.8
Breadth-across-third-unicuspids (U3B)
DSpHaOst 8.3 (0.2 3° Nie ze 0 i | Gage ate cel Ya | 655 Wea |
2.7—3.0 3.0—3.4 2.6—3.2 2.6—3.0 2.9-3.4
Breadth-across-second-molars (M2B)
5 4ee 2 oni 02 543 pmueel 0 all Soe OD 5.5. = O11
5.1-5.8 5.9-6.3 5.1-5.5 5.35.8 5.6-6.0
Palatal-length (PL)
S2ra OP? $4 02 FR pause 0 92 S50 = OZ 6.5. = U2
7.3-8.5 8.1-8.7 7.5—-8.5 8.2-8.9 8.0-8.7
Length-of-upper-toothrow (TR)
[2 a0 2 Pie farsume | E02 Pa = (Oe fess | i | dies get BS
6.6—7.5 7.4-7.8 6.8—7.4 7.2—7.5 7.2-8.0
Length-of-unicuspid-toothrow (UTR)
245-10 DAO 2A OM 2 SAO ay 2 Ot
2.1—2.5 2.42.6 2.1—2.6 2.4—2.6 2.3—2.7
Length-of-upper-molariform-toothrow (MTR)
a.2 = Ol SP et (0 Id 2 + Ol 573) = OR 54! I. 2
4.9-5.5 5.45.8 4.9-5.4 5.1-5.4 5.3-5.8
VOLUME 113, NUMBER 2 343
Table 1.—Continued.
C. mexicana
(Oaxaca) C. nelsoni C. obscura C. peregrina C. phillipsii
Posterior-width-of-M! (WM1)
Lo = 0.1 1.9 eg Onl eBrs=O:1 tc pica rg |
1.5-1.8 — 1.6-1.8 1.6—2.0 1.6—2.0
(n = 25) (n = 1)
Length-of-mandible (LM)
6.0.+ 0.2 6.4.+ 0.1 aie. 90.2 G2 20:2 64. +.0.3
5.4—6.3 6.36.6 5.4-6.4 5.86.4 5.8-7.1
Height-of-coronoid-process (HCP)
4.3 + 0.1 4.8 + 0.1 44+ 0.1 es se Oo 43 + 0.1
4.1.6 4.74.8 4.0-4.6 4.3-4.7 4.7-5.1
Height-of-coronoid-valley (HCV)
26 +. 0.1 26 = Ol Pe = (4 PS.5051 Deduct Oy]
2.32.8 2.8-2.9 2.42.8 2.7—3.0 2.72.9
Height-of-articular-condyle (HAC)
Be =*0:.2 3.9 ='0.05 So == 10:2 5.8: 0:1 GS ae
3.43.9 3.84.1 3.2-3.9 3.64.1 3.84.2
Breadth-of-articular-condyle (BAC)
mo 0.1 i Ae seee 0| 29) = Oh S30 051 cs ie aa, 6 Bs |
2.7—3.2 3.0—3.2 2.6—3.0 2.8-3.2 3.0-3.3
Articular-condyle-to-M, (AC3)
ae 30.2 a2 =) Ox A. G.==.O:2 Sb cae) 2 3.0.2. 02
4.4-5.2 5.05.3 4.3-5.0 4.8-5.3 4.8-5.3
Length-of-lower-toothrow
so O.1 6.02 0.1 Sp eee 0 Ua SQ) 2=0ek 5.9 = 02
5.4—-5.9 5.8-6.2 5.45.9 5.8—6.0 5.5-6.2
Length-of-lower-molar-row (m13)
fest /(). 1 AS 0). 4 42+ 0.1 44+ 0.1 4.4 + 0.2
4.14.5 4.44.6 4.0-4.4 4.2-4.5 4.0-4.8
(n= 6)
Length-of-m1 (m1L)
a ae OL 1.8 + 0.05 tec OL Le Ow a fae |
1.6—1.9 1.8-1.9 1.6—1.8 1.6—-1.8 1.6—2.0
Weight (g)
TAL 1D — OF S14 _ $.0)2- 14
5.0-9.5 5.0—10.0 6.5—10.0
(n = 37) (n = 23) (n = 19)
Cryptotis phillipsii appears as the most the more primitive members of the C. mex-
primitive member of the Cryptotis mexi- icana-group. Based on all but two charac-
cana-group, matching our supposition _ ters (relative tail length and relative height
based on the morphology of its forelimb. In of the coronoid process of the mandible) C.
contrast, Cryptotis peregrina is much more’ peregrina could, in fact, be considered a
derived and serves as a “‘link’’ between the member of the C. goldmani-group. How-
Cryptotis goldmani-group of shrews and ever, both characters are much more vari-
344
Table 2.—Factor loadings for the first two factor
axes from principle components analysis of Cryptotis
peregrina and Cryptotis phillipsii from Oaxaca. The
two axes accounted for 72% of the variation. Abbre-
viations as in Table 1.
Variable PC] EG2
UTR =0)1;70 0.378
PL =Orh95 0.523
IO —0.208 —0.458
MTR —0.334 SOAMOZ
TR —=(}335 0.233
M2B —0:337 33:25
PL es =O:3)/i1 0.221
CBE =O:3571 Uiz22
U1B =O;372 —0.198
U3B =0;373 —~O.Za9
factor 2
O C. peregrina
@ C phillipsii
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
able among the more primitive members of
the C. mexicana-group (Table 4) than we
previously realized (Woodman & Timm
1999). For this reason, we redefine the C.
goldmani-group below, excluding these two
characters.
Systematic Biology
“Cryptotis mexicana-group”’ Choate, 1970
We (Woodman & Timm 1999) recently
redescribed the Cryptotis mexicana-group
as small to medium-sized members of the
genus with long, dark dorsal pelage. All
members possess variably broadened fore-
feet and variably broadened and elongated
foreclaws; posterior border of the zygo-
matic plate usually positioned equal to, or
-6 -4 -2 0 2 4 6
factor 1
Fig. 1. Plot of factor scores from PCA of ten log-transformed measurements from 12 Cryptotis peregrina
and 14 Cryptotis phillipsii. Factor axis 1 represents size (Table 2); factor loadings are negatively weighted, so
the largest individuals have the smallest scores. Factor axis 2 represents a combination of palatal-length, inter-
orbital-breadth, unicuspid-toothrow-length, and breadth-across-M?s (Table 2).
VOLUME 113, NUMBER 2
20
1
345
C. parva
C. nigrescens
C. mexicana
E C. nelsoni
. peregrina
. alticola
C. goldmani
C. goodwini
. gnseoventris
C. obscura
C. phillipsii
Fig. 2. Topology of the single most parsimonious tree resulting from phylogenetic analysis of the Cryptotis
mexicana-group of shrews. Cryptotis nigrescens and C. parva parva served as the outgroups for polarizing
t—)
characters. Number of transitions is shown for each branch. Tree length = 67 (minimum possible length = 49;
maximum possible length = 122); consistency index =
index = 0.753; homoplasy index = 0.269.
slightly posterior to, the posterior base of
the maxillary process; upper toothrow un-
crowded; dentition not bulbous; anterior
border of the coronoid process of the man-
dible joins the horizontal ramus at a rela-
tively low angle; posterior border of lower
incisor extends to posterior cingulum of
P,; relatively long distance from the coro-
noid process to the posterior border of M;;
tall, wide articular face of the articular pro-
cess; deep lower sigmoid notch; relatively
long, low P,; relatively short, broad meta-
carpals; shortened and broadened humerus
with elongated processes and a dorsoven-
trally elongate head; deeply pocketed pos-
terior edge of the falciform process of the
tibia.
Included species.—Cryptotis alticola, C.
goldmani, C. goodwini, C. griseoventris, C.
0.731; rescaled consistency index = 0.551; retention
mexicana, C. nelsoni, C. obscura, C. pere-
grina, and C. phillipsii.
Cryptotis peregrina (Merriam, 1895)
Blarina mexicana peregrina Merriam,
1895:24.
C[ryptotis]. mexicana peregrina: Miller,
911222.
Cryptotis mexicana peregrina: Hall & Kel-
son, 1959:60; Goodwin, 1969:39 (in
part); Choate, 1970:237 (in part); Hall,
1981:59 (in part).
Cryptotis mexicana mexicana: Goodwin,
1969:39 (in part).
Cryptotis peregrina: Woodman & Timm,
1999:35 (in part).
Holotype.—Skin and skull of male,
USNM 68317; captured on 12 September
346
Table 3.—Complete character matrix for phylogenetic analysis of members of the Cryptotis mexicana-group and two outgroups, Cryptotis parva parva, and Cryptotis
nigrescens. Transition series (TS) are explained in Appendix 1.
Transition series
N
Taxon
parva
nigrescens
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Sc A ce re ce ee ee ee en |
SKS SES iw i i i oo
See eee ee ee!
N
mexicana
N
nelsoni
obscura
peregrina
phillipsii
alticola
goldmani
[oe)
goodwini
3
griseoventris
1894 by E. W. Nelson and E. A. Goldman
(collector number 6748).
Type locality.—‘‘mountains 15 miles
[south]west of city of Oaxaca [de Juarez],
Mexico (altitude, 9,500 feet).’’ According
to Goldman (1951), the collectors spent
September 10th to 19th, 1894, traveling to
and from, and working at, this locality. This
trip took them ca. 8 mi southwest of Oaxaca
de Juarez to Cuilapan, then west into the
mountains, passing Santa Inéz [Santa Inés
del Monte, 16°54’30"N, 96°52’W] and
climbing over the summit to a wet meadow
called Neverfa Herrera on the western
slope.
“These high mountains, which appeared to have no
distinctive name, are somewhat isolated, the only
connection with the high mountains to the east be-
ing through low hills north of Oaxaca. The upper
slopes are well watered, and general conditions
much as on Cerro San Felipe. Specimens were la-
beled “Mountains 15 miles west of Oaxaca,’ but as
we later learned the direction was more nearly
southwest.”’ (Goldman, 1951:218).
Merriam (1895) reported that the speci-
mens were obtained from a variety of ele-
vations between 8800 [2680 m] and the
summit at 9500 ft [2895 m], indicating that
the type series of Cryptotis peregrina was
collected at or near the top of the range. On
topographic maps, the summit appears to be
near a landmark called La Plazuela
[16°56'52”N, 96°53'45"W]. Binford (1989)
refers to these mountains as the Sierra de
Cuatro Venados.
Distribution.—Known from 2680 to
3200 m in the Sierra de Cuatro Venados,
central Oaxaca, and the Sierra Yucuyacua,
western Oaxaca (Fig. 3).
Description.—Size medium to large for
the genus (Table 1); tail long, averaging 30
mm, or ca. 42% (Table 4) of head-and-body
length; dorsal guard hairs 6—7 mm long;
forepaws enlarged; foreclaws noticeably
elongate and broadened; rostrum of mod-
erate length (PL/CBL = 44.5%, Table 4);
often two well-developed dorsal foramina
(44%, Table 4); foramen posterior to dorsal
articular facet leading to ventral extension
VOLUME 113, NUMBER 2
Oaxaca
re de Juarez
Fig. 3.
of the sinus canal typically present on one
or both sides of the cranium (94%, Table
4); foramen dorsal to dorsal articular facet
typically absent (87%, Table 4); zygomatic
plate narrow (ZP/PL = 19.3%, Table 4), an-
terior border usually aligned with posterior
¥%3 of mesostyle-metastyle valley or metas-
tyle of M'; U* unreduced in size, surface
area >'% that of U’; U* usually aligned with
the unicuspid toothrow and partially visible
in lateral view of the skull; P*, M', and M?
slightly to moderately recessed on posterior
border; protoconal basin of M! reduced rel-
347
Map of part of Oaxaca, Mexico, illustrating the known distribution of Cryptotis peregrina (*) and
Cryptotis phillipsii (@). The 2000 m contour is shown.
ative to hypoconal basin; M? simple: usu-
ally possessing paracrista, paracone, pre-
centrocrista (all pigmented), mesostyle, and
very short postcentrocrista; M?* lacking
metacone, hypocone absent or poorly-de-
veloped, and a reduced, typically unpig-
mented protocone occasionally present. Ar-
ticular process of the mandible generally
moderately tall and wide, with a moderately
broad lower articular facet; moderately high
coronoid process of the mandible (HCP/ML
= 73.1%, Table 4); entoconid usually pre-
sent on M, (73%, Table 4), typically vesti-
348 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 4.—Characteristics among the five species previously synonomized with Cryptotis mexicana.
C. mexicana
(Oaxaca) C. nelsoni C. obscura C. peregrina C. phillipsii
Foramen of sinus canal
7% O% 19% 94% 24%
n = 90 n=8 n = 52 n= 16 n= 17
Foramen dorsal to dorsal articular facet present
13% 38% 28% 13% 82%
n = 90 n=8 n = 36 n= 15 n=17
Two distinct dorsal foramina present
54% 50% 22% 44% 47%
n = 95 n=6 n= 58 n= 16 n= 19
Posteroventral border of unicuspids
concave concave straight to convex concave concave
Entoconid of M, present
100% 100% 96% 73% 93%
68% well-developed 100% well-developed 96% well-developed 100% vestigal 83% vestigal
= 77 n=7 n = 26 n = 22 n= 13
Humerus modified
slightly slightly extremely slightly
Relative tail length (TL/HB x 100)
392 5 38 + 4 40° + 6 42 iS 46 + 6
26-52 28-44 29-55 36-47 30-59
n= 118 n=9 n= 55 n = 24 n = 27
Relative rostrum length (PL/CBL xX 100)
Asal = 0:8 45.2, = 0.8 ya i use [| 44.5.+ 0.7 433 = 0.6
41.7-45.1 42.0-44.3 41.0-45.6 43.246.0 42.244.4
n = 30 n=6 n = 32 n= 12 n= 14
Relative breadth of zygomatic plate (ZP/PL x 100)
2166: = V6 23a A PAST ae bal 19.32 .1.0 234° 2-46
17.9-24.4 21.8—25.6 22.2—26.9 18.1-21.2 21.4—26.4
n = 30 n=7 n = 32 n= 12 n= 14
Relative breadth of zygomatic plate (ZP/CBL Xx 100)
9.4 + 0:7 10:2 = 0:6 10:6: ="65 S67 0'5 LO = Os
7.9-10.4 9.5—11.2 9.7-11.5 7.8—-9.4 9.2-11.4
n = 30 n=6 n = 32 n= 12 n= 14
Relative length of unicuspid toothrow (UTR/CBL X 100)
12a) 0.4 12 Duct 0,5 13:0) 27054 129° +03 12 EO:6
11.4-13.3 12.0—-13.4 11.6—13.9 12.1-13.4 11.4—-13.7
n = 30 n=6 n = 32 n= 12 n= 14
Relative palatal breadth (M2B/PL Xx 100)
65:9 = 2.5 T2o = OO 67.4 + 2:2 63.2 se DS 6827 271
60.0—72.6 70.9-73.3 62.4—70.7 60.9—-68.7 65.1—72.5
n = 30 n=7 n = 32 n= 12 n= 14
Relative height of coronoid process (HCP/ML x 100)
TOS = ScD 73.8 13 74.3-= 2:0 MSA es 16 Tse) tae
65.1—79.6 72.3-76.2 70.2—83.6 69.4—75.0 71.6—86.2
n = 30 n=8 n = 32 n= 12 n= 16
VOLUME 113, NUMBER 2 349
Table 4.—Continued.
C. mexicana
(Oaxaca) C. nelsoni C. obscura C. peregrina C. phillipsii
Relative posterior length of mandible (AC3/ML X 100)
80.8 + 2.6 yA aos a 77.0 + 3.0 Seal = yh 1484 = 3.6
77.4—86.2 77.8-82.5 72.6—85.5 77.8-85.0 73.2-86.2
n = 30 n= 8 n = 32 n= 12 n= 16
Relative extension of articular condyle (AC3/HCP x 100)
Haw = 5.) 108:8-+ 1,7 105.6) 37 iA 20 103.7 = 3.9
100.0—124.4 106.4—110.6 100.0—114.3 109.9-115.6 96.1—110.6
n = 30 n= 8 n = 32 n= 12 n= 16
gial and unpigmented when present. Broad,
curved humerus, with elongated processes
(Fig. 4B).
Comparisons.—Cryptotis peregrina is
distinctive among the species previously
considered to be subspecies of Cryptotis
mexicana. It has a narrow zygomatic plate,
and it has the broadest forepaws and the
longest and broadest foreclaws. The hu-
merus is shorter, more curved, and gener-
ally more highly derived: the pectoral pro-
cess is higher, lateral epicondyle more ex-
Fig. 4 Left humeri of (A) Cryptotis phillipsii (KU 124299) and (B) Cryptotis peregrina (AMNH 149965).
The humerus labeled “C. peregrina” in Woodman and Timm (1999: Fig. 15B) is from a specimen (KU 124298)
that we now recognize as C. phillipsii.
350
panded, and teres tubercle and medial epi-
condyle longer and more closely situated to
each other. Cryptotis peregrina is the only
species in the C. mexicana-group besides
Cryptotis goldmani that usually has a well-
developed foramen of the ventral branch of
the sinus canal (Table 4; Woodman &
Timm 1999).
Cryptotis mexicana: Cryptotis peregrina
has slightly longer, broader claws and a
much more derived humerus; M®? simple,
lacking metacone; entoconid of M, vesti-
gial (rather than well-developed), when
present.
Cryptotis nelsoni: Cryptotis peregrina
has slightly longer, broader claws and a
much more derived humerus; skull narrow-
er overall; longer rostrum and much nar-
rower palate (Table 4); M? simple, lacking
metacone; entoconid of M, vestigial (rather
than well-developed), when present.
Cryptotis obscura: Cryptotis peregrina
has much longer, broader claws and a much
more derived humerus; longer rostrum (Ta-
ble 4); upper unicuspids concave on pos-
terior surface (rather than straight or con-
vex), appearing curved; M> simple, lacking
metacone; more likely to have two well-de-
veloped dorsal foramina (44% vs. 22%, Ta-
ble 4); entoconid of M, vestigial (rather
than well-developed), when present.
Cryptotis phillipsii: Cryptotis peregrina
has much longer and broader claws and a
much more derived humerus; longer ros-
trum and much narrower palate (Table 4);
absolutely and relatively broader zygomatic
plate (Tables 1, 4); larger U* relative to U’;
slightly more recessed upper dentition;
slightly less complex M?; well-developed
foramen of the ventral extension of sinus
canal; usually no foramen dorsal to dorsal
articular facet.
Cryptotis goldmani-group: Cryptotis per-
egrina is generally smaller in head-and-
body-length and has a relatively longer tail
(Table 1, 4); narrower zygomatic plate;
more likely to possess a vestigial entoconid
(rather than no entoconid) of M?.
Remarks.—Nearly all known specimens
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
of Cryptotis peregrina are part of the type
series, collected in the Sierra de Cuatro
Venados ca. 15 mi SW of Oaxaca de Jua-
rez in 1894. Merriam (1895) stated that
there were 25 specimens in the type series,
but he did not list them individually. We
were able to locate only 24 original spec-
imens collected by Nelson and Goldman.
The only other specimen of C. peregrina
of which we are aware is a fluid-preserved
individual (with the skull and left humerus
removed) in poor condition, collected “‘N.
La Muralla”’ in the Sierra Yucuyacua by
Thomas B. MacDougall on 18 March
1952. Locality notes by MacDougall indi-
cate this locality is at 10,500+ ft at the
‘“‘top of Cerro Yucunino” (AMNH field
notes, 1952). We believe (as apparently did
Choate, 1970:239) that this peak is equiv-
alent to Cerro Yucuyacua [= Piedra de
Ollila], based on its elevation and position
relative to La Muralla and Santa Maria
Asuncion Tlaxiaco.
Goldman (1951:218) noted that the type
locality was moist, with general conditions
‘“‘much as on Cerro San Felipe.’’ Areas
above 8000 ft on Cerro San Felipe were
covered in oak-dominated mixed forest
with an herbaceous understory, and they
experienced cold night-time temperatures,
with frosts recorded in August. Meadows
(neverias), such as the one in the vicinity
of the type locality, were said to be em-
ployed by local people for making ice to be
used at lower elevations. MacDougall
(AMNH field notes, 1952) noted the pres-
ence of pine and oak where he captured C.
peregrina north of La Muralla.
Specimens examined (25).—Mexico: Oa-
xaca: Nforth]. [of] La Muralla [16°58'N,
97°56'W], top of Cerro Yucunin6 [= Cerro
Yucuyacua, 17°06’N, 97°40'W], Tlaxiaco
[Santa Maria Asuncion Tlaxiaco, 17°16'N,
97°41'W], 10,500+ ft (AMNH 149965);
mountains 15 mi [south] west Oaxaca de
Judrez [16°30°N, 96°33 Wi. 9500 aa
(BMNH 95.11.1.26 [= USNM 68337],
USNM 68315-68336, 68338, including ho-
lotype).
VOLUME 113, NUMBER 2
‘““Cryptotis goldmani-group’’> Woodman &
Timm, 1999
Description.—The C. goldmani-group is
a subset of the C. mexicana-group that is
comprised of medium-sized members of the
genus with greatly broadened forefeet; ex-
tremely long, broad foreclaws; fourth upper
unicuspid usually aligned with the unicus-
pid toothrow and partially visible in labial
view of the cranium; protoconal basin of
M! reduced relative to hypoconal basin; M°
simple, hypocone absent or poorly devel-
oped and lacking metacone; entoconid of
M, vestigial or absent; and extremely broad
humerus with greatly elongated processes.
Included species.—Cryptotis alticola, C.
goldmani, C. goodwini, C. griseoventris,
and C. peregrina.
Cryptotis phillipsii (Schaldach, 1966)
Notiosorex (Xenosorex) Phillipsii Schal-
dach, 1966:289; Goodwin, 1969:43.
Cryptotis mexicana machetes: Musser,
1964:6; Schaldach, 1966:288; Goodwin,
1969:40 (in part).
Cryptotis mexicana peregrina: Goodwin,
1969:40 (in part); Choate, 1970:237 (Gn
part); Hall, 1981:59 (in part).
Cryptotis peregrina: Woodman & Timm,
i225) (in part).
Holotype.—Skin and skull of adult fe-
male, UNAM 8445; taken 18 Dec 1964 by
W. J. Schaldach, Jr. (collector number
13278). Two paratypes, KU 114226 [=
UNAM 8446] and UNAM 8447.
Type locality.—*‘the Rio Molino, 3 ki-
lometers S.W. San Miguel Suchixtepec, al-
titude 2250 meters, southern Oaxaca, Méx-
ico”’ (Schaldach, 1966:289).
Distribution.—Known from 1060 to
2745 m in the Sierra de Miahuatlan, south-
ern Oaxaca, Mexico (Fig. 3).
Description.—Size medium to large for
the genus (Table 1); tail long, averaging 31
mm, or ca. 46% (Table 4) of head-and-body
length; dorsal guard hairs 6-7 mm long;
forepaws somewhat enlarged; foreclaws
351
among the narrowest for members of the
Cryptotis mexicana-group, but elongate and
somewhat broadened relative to other mem-
bers of the genus; rostrum of moderate
length (PL/CBL = 43.3%, Table 4); often
two well-developed dorsal foramina (47%,
Table 4); ventral extension of the sinus ca-
nal and associated foramen typically lack-
ing (76%, Table 4), but well-developed
when present; a foramen dorsal to the dor-
sal articular facet usually present on one or
both sides of the skull (82%, Table 4); zy-
gomatic plate broad (ZP/PL = 23.4%, Table
4), anterior border usually aligned with pos-
terior % of mesostyle-metastyle valley or
metastyle of M'; P*, M', and M? unrecessed
to slightly recessed on posterior border; sur-
face area of U* typically =% that of U*; U*
usually aligned with the unicuspid tooth-
row, but not typically visible in lateral view
of the skull; protoconal basin of M' same
size as hypoconal basin; M? simple, lacking
metacone, hypocone poorly-developed or
absent; M? usually possessing paracrista,
paracone, precentrocrista, and mesostyle
(all pigmented), and occasionally a very
short postcentrocrista and a well-developed,
pigmented protocone. Articular process of
the mandible generally moderately tall and
wide, with a moderately broad lower artic-
ular facet; coronoid process of the mandible
moderately high relative to mandibular
length (HCP/ML = 75.7%, Table 4); ento-
conid usually present on M, (92%, Table 4),
typically vestigial (83% of those possessing
entoconids), but occasionally well-devel-
oped (17%). The tall, broad, slightly curved
humerus has somewhat elongated process-
es, but it is among the least modified within
the C. mexicana-group (Fig. 4).
Comparisons.—Cryptotis phillipsii has
the least modified forelimbs of any member
of the Cryptotis mexicana-group: it has the
smallest forepaws and the shortest and nar-
rowest foreclaws; the humerus is among the
longest and least curved; teres tubercle is
short, medial epicondyle is short and has a
Straight proximal edge, and teres tubercle
and medial epicondyle are broadly separat-
352
ed. Cryptotis phillipsii also has the longest
tail relative to head-and-body-length (Table
4). There may be a local tendency for some
C. phillipsii to develop without U*s (12%,
n = 17) in the region of the type locality,
resulting in a reduced dentition.
Cryptotis mexicana: Cryptotis phillipsii
has shorter, narrower foreclaws and a rela-
tively longer tail; less modified humerus;
broader palate (Table 4); simple M?, lacking
metacone; entoconid of M, usually vestigial
(rather than well-developed) when present.
Cryptotis nelsoni: Cryptotis phillipsii has
shorter, narrower foreclaws and a relatively
longer tail; narrower palate (Table 4); sim-
ple M?, lacking metacone; entoconid of M,
usually vestigial (rather than well-devel-
oped) when present.
Cryptotis obscura: Cryptotis phillipsii
has a relatively longer tail; upper unicus-
pids concave on posterior surface, appear-
ing curved, rather than convex and cone-
shaped; more likely to have two well-de-
veloped dorsal foramina (Table 4); simple
M?, lacking metacone; entoconid of M, usu-
ally vestigial (rather than well-developed)
when present.
Cryptotis goldmani-group: Cryptotis
phillipsii is smaller in head-and-body-
length, and has much shorter, narrower fo-
reclaws, a much less derived humerus, and
a relatively longer tail; higher coronoid pro-
cess (Table 4); shorter posterior portion of
mandible (Table 4); much more likely to
possess vestigial entoconid (rather than no
entoconid) of M,.
Remarks.—The type series of Cryptotis
phillipsii was collected in 1964 by William
J. Schaldach, Jr. and Allan R. Phillips dur-
ing a trip to southern Oaxaca to obtain
specimens of birds and mammals. Schal-
dach (1966) originally described a new spe-
cies, Notiosorex phillipsii, on the basis of
three specimens: a skin with skull and a
skin without skull from Rio Molino
(UNAM 4445, 4447) and a skin with skull
from Rio Guajalote (KU 114226 = UNAM
8446). Originally identified as Cryptotis
mexicana, the main consideration in sub-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
sequently referring these three specimens to
Notiosorex appears to have been the pres-
ence of only three unicuspids in the upper
toothrow, rather than the four unicuspids
typical of Cryptotis. A fourth specimen
(UNAM 8444), collected at Rio Molino and
possessing four unicuspids, was reported as
Cryptotis mexicana (Schaldach 1966). Not-
ing strong differences in external, cranial,
and dental characters between the type se-
ries of N. phillipsii and specimens of Notio-
sorex crawfordi and Megasorex gigas that
he used for comparison, Schaldach (1966)
erected a new subgenus, Xenosorex, to dis-
tinguish the new shrew. Choate (1969) re-
examined the type series of Notiosorex
phillipsii and reported a number of distinc-
tive cranial, mandibular, and dental char-
acters that clearly established N. phillipsii
as a member of the genus Crypfotis, despite
its reduced dentition. He justifiably placed
Schaldach’s N. phillipsii as a junior syno-
nym of Cryptotis [mexicana] peregrina,
where it has remained until now (Choate
1970, Hall 1981, Hutterer 1993, Woodman
& Timm 1999).
Cryptotis phillipsii is known from higher
elevation forest, including cloud forest, in
the Sierra de Miahuatlan of southern Oa-
xaca. The species is syntopic with Cryptotis
goldmani at some localities, but the geo-
graphic distribution of C. phillipsii is not
nearly as broad as that of C. goldmani, de-
spite an apparently lower elevational limit.
Schaldach (1966) reported the humid,
shaded ravines along the slopes of the val-
ley of the Rio Molino at the type locality
contained remnant, pine- and oak-dominat-
ed cloud forest, with a dense understory of
ferns, mosses, vines, and shrubs. Musser
(1964) described the canyon bottom as con-
taining moist, open forest with bromeliad-
covered oaks and a dense ground cover of
shrubs, ferns, and herbaceous vegetation.
The higher, drier slopes were covered with
a mixed secondary growth of pines and
oaks. Musser (1964:6) took one C. phillip-
sil, two Sorex saussurei oaxacae, and 29
Peromyscus megalops under a “‘lush her-
VOLUME 113, NUMBER 2
baceous ground cover’’ consisting largely
of ferns along the steep banks of a “‘moist,
densely vegetated streamside”’ with a “‘deep
layer of wet humus and leaf litter’? at 7300
ft [2225 m] near Rio Molino. Paul B. Rob-
ertson (KU field notes and catalog, 1969,
1970) captured three C. phillipsii at Rio
Molino. In December 1969, he trapped one
C. phillipsii, one Microtus mexicanus, two
Peromyscus aztecus, two P. levipes, and
four P. megalops. In April 1970 using a
trapline “‘along a small rivulet,’ he took
one C. phillipsii with two Cryptotis gold-
mani, one Sorex saussurei oaxacae, one
Microtus mexicanus, one Neotoma mexi-
cana, two Oryzomys chapmani, 12 Pero-
myscus megalops, and one Reithrodonto-
mys mexicanus. Nearby, “‘among & under
rocks in an overgrown field,’’ he collected
a C. phillipsii with two Reithrodontomys
megalotis, three R. sumichrasti, and one
Sigmodon alleni. He described the stream-
side vegetation at Rio Molino as “hard
wood”’ and the slopes as “almost pure
pine.’ In addition, Schaldach (1966) re-
ported capturing Sorex mutabilis (as Sorex
veraepacis) and Liomys pictus at Rio Mol-
ino. Cryptotis phillipsii and C. goldmani
were collected in syntopy also at “‘lumber
camp, km 158” along the Puerto Angel
Road in January 1970 by John R. Arnold
and Craig Moe. Cryptotis phillipsii and C.
parva pueblensis were captured syntopical-
ly 20 mi S, 5 mi E San Miguel Sola de Vega
by Percy L. Clifton in June—July 1964. He
described the habitat there as dense cloud
forest, with C. phillipsii “‘caught under a
rock in a low damp situation along a creek”’
(KU field notes, 1964). Cryptotis phillipsii
also was taken in syntopy with C. p. pue-
blensis at San Miguel Suchixtepec by
Thomas B. MacDougall in October 1967.
Specimens examined (36).—Mexico: Oa-
xaca: 20 mi S, 5 mi E Sola de Vega [San
Miguel Sola de Vega, 16°31'N, 96°59’W],
4800 ft (KU 98728); 36 km (by road) N
San Gabriel Mixtepec [16°06’N, 97°06'’W],
1680 m (KU 124294); 27.8 km (by road)
N San Gabriel Mixtepec, 1320 m (KU
353
124295, 124296); La Cima, Puerto Escon-
dido road, km 184.5 [16°12'N, 97°07'W],
5750 ft (CAS 15473); Puerto Escondido
road, km 193 [16°10’N, 97°07'W], 4200 ft
(CAS 15474); Sinai [Finca Sinai, 16°07'N,
97°08'W], 10 km [by trail] E Nopala [Santo
Reyes Nopala], 7200 ft (CAS 14940); Rio
Molino [16°04'N, 96°28'’W], 2250-2745 m
(AMNH 213758, 213759, 214152, 214803-
214805; KU 121661, 124298, 124299;
UNAM 8444, 8445, 8447—includes holo-
type); near campemento Rio Molino (Hwy
175), 7300 ft (UMMZ 112572); Puerto An-
gel road, km 153 [16°04'N, 96°28’W], 7100
ft (CAS 14068); San Miguel Suchixtepec
[16°OS’N, 96°28’W], Miahuatlan District
(AMNH 214806—214808); 16 km SW [San
Miguel] Suchixtepec, 2000 m (ENCB 3413,
3414); Lovene [16°02’N, 96°12’W], Mia-
huatlan (AMNH_ 178739); lumber camp,
Puerto Angel road, km 158, 8375 ft (CAS
15478); Rio Guajalote [16°00'N, 96°28’ W],
2000 m (KU 114226); Rio Jalatengo
[15°58’N, 96°27'W], Puerto Angel road, km
178, 4275 ft (CAS 14069, 14071, 14072,
15475; UNAM 27518); Puerto Angel road,
km 195, 3475 ft (UNAM 26551); Puerto
Angel road (UNAM 27517).
Acknowledgments
We thank Robert P. Anderson for valu-
able assistance in tracking down informa-
tion helpful to our study. We thank the fol-
lowing curators and collection managers for
loans and for permission to study important
specimens under their care: Guy G. Musser
and Robert Voss (AMNH); Robert C. Dow-
ler (ASNHOC); Paula Jenkins (BM); Douglas
J. Long (CAS); Ticul Alvarez (ENCB);
John E. Heyning (LACM); Maria E. Rutz-
moser (MCZ); Elmer C. Birney (MMNH);
Livia Leén P. (MZFC); George D. Baum-
gardner (TCWC); Phil Myers (UMMZ);
Fernando Cervantes R. (UNAM); Michael
D. Carleton, Alfred L. Gardner, Linda K.
Gordon, Helen L. Kafka, and Richard W.
Thorington, Jr. (USNM). Two reviewers
354
provided helpful comments that improved
our manuscript.
Literature Cited
Binford, L. C. 1989. A distributional survey of the
birds of the Mexican State of Oaxaca.—Orni-
thological Monographs 43:1—418.
Choate, J. R. 1969. Taxonomic status of the shrew,
Notiosorex (Xenosorex) phillipsii Schaldach,
1966 (Mammalia: Insectivora).—Proceedings
of the Biological Society of Washington 82:
469-476.
. 1970. Systematics and zoogeography of Mid-
dle American shrews of the genus Cryptotis.—
University of Kansas Publications, Museum of
Natural History 19:195-—317.
Goldman, E. A. 1951. Biological investigations in
México.—Smithsonian Miscellaneous Collec-
tions 115:1—xiv, 1—476.
Goodwin, G. G. 1969. Mammals from the state of Oa-
xaca, Mexico, in the American Museum of Nat-
ural History.—Bulletin of the American Muse-
um of Natural History 141:1—270.
Hall, E. R. 1981. The mammals of North America, 2nd
edition. John Wiley & Sons, New York, 1:1—
600 + 90 pp.
, & K. R. Kelson. 1959. The mammals of North
America. The Ronald Press Company, New
York, 1:1-546 + 79 pp.
Maddison, W. P., & D. R. Maddison. 1992. MacClade:
analysis of phylogeny and character evolution,
version 3. Sinauer Associates, Sunderland, Mas-
sachusetts.
Merriam, C. H. 1895. Revision of the shrews of the
American genera Blarina and Notiosorex.—
North American Fauna 10:5—34.
Miller, G. S., Jr. 1911. Three new shrews of the genus
Cryptotis.—Proceedings of the Biological So-
ciety of Washington 24:221—224.
Musser, G. G. 1964. Notes on geographic distribution,
habitat, and taxonomy of some Mexican mam-
mals.—Occasional Papers of the Museum of
Zoology, University of Michigan 636:1—22.
Reed, C. A. 1951. Locomotion and appendicular anat-
omy in three soricoid insectivores.—American
Midland Naturalist 45:513—671.
Schaldach, W. J., Jr. 1966. New forms of mammals
from Southern Oaxaca, Mexico, with notes on
some mammals of the coastal range.—Sauge-
tierkundliche Mitteilungen 4:286—297.
Swofford, D. L. 1998. PAUP*. Phylogenetic analysis
using parsimony (*and other methods). Version
4. Sinauer Associates, Sunderland, Massachu-
setts.
Woodman, N., & R. M. Timm. 1992. A new species
of small-eared shrew, genus Cryptotis (Insectiv-
ora: Soricidae), from Honduras.—Proceedings
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
of the Biological Society of Washington 105:1—
12
5 OE . 1993. Intraspecific and interspe-
cific variation in the Cryptotis nigrescens spe-
cies complex of small-eared shrews (Insectivo-
ra: Soricidae), with the description of a new
species from Colombia.—Fieldiana: Zoology
(New Series) 74:1—30.
SEGG . 1999. Geographic variation and
evolutionary relationships among broad-clawed
shrews of the Cryptotis goldmani-group (Mam-
malia: Insectivora: Soricidae).—Fieldiana: Zo-
ology (New Series) 91:1—35.
Appendix I: Transition Series Used in Phylogenetic
Analysis
Transition series (TS) marked by an asterisk (*)
were modified from Woodman & Timm (1999). TS
30—32 were added since that paper.
*1. length of foreclaws: short (0); elon-
gate (1); more elongate (2); greatly
elongate (3).
*2. breadth of foreclaws: narrow (0);
broad (1); broader (2); extremely
broad (3).
3. forefeet: small (0); enlarged, broad-
ened (1); greatly enlarged and broad-
eneau(2).
4. metacarpals: long, narrow (0); short,
broad (1).
5. posterior border of zygomatic plate:
even with or anterior to anterior root
of maxillary process (0); even with
posterior root of zygomatic process,
but separated from it by posterior bor-
der of palate (1); even with (or pos-
terior to) and confluent with posterior
root of zygomatic process (2).
6. anterior border of coronoid process:
steep, forming a narrow angle with
horizontal ramus of mandible (Q); less
steep, forming a wide angle with hor-
izontal ramus of mandible (1).
7. articular condyle: low and broad (0);
high and narrow (1).
8. lower sigmoid notch: very shallow
(O};-deep’ (Pf):
9. shape of unicuspids (U!—U*): cone-
shaped, posteroventral border straight-
edged or convex (0); narrow, poster-
oventral border concave (1).
10. protoconal basin of M!: about equal in
VOLUME 113, NUMBER 2
i.
ee
1S.
14.
15.
*16,
eat.
ca >
£S:
20.
DA.
cpp 24
size to hypoconal basin (0); reduced
relative to hypoconal basin (1).
M? morphology: simple, metacone ab-
sent (O); complex, metacone present
Ch).
shape of P,: short and high (0); long
and low (1).
shape of humerus: long, narrow, rela-
tively straight (0); short, robust,
curved (1); short, robust, and very
curved (2).
head of humerus: rounded (0); dorso-
ventrally elongate (1).
ventral edge of proximal face of great-
er tuberosity of humerus: rounded (0);
with broad, deep pocket (1).
pectoral process of humerus: low (0);
high (1).
length of teres tubercle: relatively
short (0); elongate (1); greatly elon-
gate (2).
medial epicondyle of humerus: short
(O); elongate (1); greatly elongate (2).
lateral epicondyle of humerus: small
(O); expanded (1).
teres tubercle and medial epicondyle
of humerus: far apart (0); close to-
gether (1); very close (2); extremely
close (3).
posterior edge of falciform process of
tibia: not deeply pocketed (0); deeply
pocketed (1).
foramen of sinus canal: absent in
100% of specimens (0); absent in
>75% of specimens (1); present in
>50% of specimens, but vestigial (2);
present in >90% of specimens, well-
developed (3).
24.
F202;
26.
Dike
28.
Pas J
#30;
pele
*52.
. dorsal foramina:
two in <75% of
specimens (0); two in >75% of spec-
imens (1).
foramen dorsal to articular facet: pre-
sent in <75% of specimens (0); pre-
sent in >75% of specimens (1).
entoconid of M,: present in >90% of
specimens, typically well-developed
(O); present in >50% of specimens,
typically vestigial (1); absent in >80%
of specimens (2).
body size (head-and-body length):
smaller, mean <69 (QO); larger, mean
> ie),
relative tail length (% of head-and-
body length): short, mean <39% (0);
long, mean >40% (1).
upper unicuspid toothrow: crowded,
three unicuspids visible in lateral view
(0); uncrowded, four unicuspids visi-
ble in lateral view (1).
dorsal guard hairs: short, <4 mm (0);
of intermediate length, 4-5 mm (1);
long, >5 mm (2).
distal edge of pectoral process of hu-
merus relative to teres tubercle: ex-
tends well distal to teres tubercle (0);
extends to proximity of distal edge of
teres tubercle (1).
bones of hind paws: metatarsals and
proximal phalanges long narrow (0);
metatarsals and proximal phalanges
broad (1); metatarsals and proximal
phalanges short and very broad (2).
posterior border of P*, M', and M7’:
slight to no emargination (0); notica-
ble emargination (1); strong emargi-
nation (2).
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):356—368. 2000.
Two new species of flightless rails (Aves: Rallidae) from the Middle
Pleistocene ‘‘crane fauna”? of Bermuda
Storrs L. Olson and David B. Wingate
(SLO) Department of Vertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560-0116, U.S.A.;
(DBW) Bermuda Natural History Museum, P.O. Box FL 145, Flatts, Bermuda
Abstract.—Two new species of flightless rails are described from a Pleis-
tocene fauna in Bermuda that also includes an extinct crane (Grus latipes) and
an extinct duck (Anas pachyscelus). The medium-sized Rallus ibycus, new
species, was possibly derived from North American populations of Virginia
Rail (R. limicola), but had a longer bill, much more robust legs, and reduced
wings and pectoral girdle. The very small Porzana piercei, new species, except
for the reduced wing and pectoral girdle, is very similar to the extant Yellow-
breasted Crake (P. flaviventer), which now occurs only in the Neotropics, in-
cluding the Greater Antilles. The fauna that included these rails developed
during a long, stable glacial period of lowered sea-levels in the Middle Pleis-
tocene, during which the entire Bermuda platform was emergent. This was
followed by an abrupt and extreme interglacial about 400,000 years ago when
sea-levels rose to 21 m above present levels, obliterating most of Bermuda and
much of its endemic fauna, including the rails.
The island of Bermuda, situated in the
western North Atlantic 1050 km east of
Cape Hatteras, North Carolina, is composed
almost entirely of calcareous aeolianite on
the southeastern rim of a submerged, trun-
cated summit of an extinct volcano. The
aeolianite has been modified by solution to
form numerous caves and fissures that are
accessible to collectors either through nat-
ural openings or through limestone quar-
rying operations. These caves frequently
contain accumulated fossils of vertebrates
and invertebrates, mainly birds and terres-
trial gastropods. Although much of interest
to avian paleontology has been collected
from Bermuda, relatively little has been
published, partly for lack of a better under-
standing of the complexities of the island’s
stratigraphy and chronology, which has
been greatly improved in recent years.
The first contribution to knowledge of
Pleistocene birds in Bermuda was that of
Wetmore (1960) who described a new ge-
nus and species of endemic crane, Baeo-
pteryx latipes, and an extinct endemic duck,
Anas pachyscelus, from the Wilkinson
Quarry, Hamilton Parish. The genus Baeo-
pteryx was later synonymized with typical
cranes of the genus Grus (Fischer & Ste-
phan 1971).
Wetmore (1960:10) mentioned that “‘the
collection contains various bones from four
species of rails, one very small, two of in-
termediate size, and one nearly as large as
the modern clapper rail [Rallus longiros-
tris]. These are not clearly marked in the
present collection so that no attempt is
made to describe them here in detail, par-
ticularly since complete material for one of
them is now in other hands for study.’’ Ac-
tually, among the rail material that Wetmore
examined, there are remains of only two
species of rails, one medium-sized and the
other very small (uncataloged specimens in
VOLUME 113, NUMBER 2
USNM). Wetmore’s information on Ber-
muda rails apparently came in part from
Pierce Brodkorb, University of Florida,
who had been in Bermuda collecting fossil
rails with Wingate only weeks before Wet-
more’s publication appeared. Later, Olson
(1977:353-354) briefly mentioned the still
undescribed fossil rails from Bermuda.
Brodkorb’s collections containing the Ber-
muda rails passed to the Florida Museum
of Natural History after Brodkorb’s death
in 1992, and we have now belatedly begun
the process of describing them.
Some information on the relationships
and adaptations of the largest species of rail
to which Wetmore alluded was provided by
Olson (1997). This species was not contem-
poraneous with the “‘crane fauna’”’ and does
not appear to have co-existed with any oth-
er endemic rail on Bermuda. In February
1999, we discovered a new deposit in
which this species was the predominant
bird. Preliminary results from amino acid
racemization ratios from associated snail
shells indicate that it originated during a
much younger glacial period. We have post-
poned description of this species until the
new material can be fully prepared and an-
alyzed. Here we describe the two species
that are definitely part of the “‘crane fauna”’
that we believe dates back at least to the
Middle Pleistocene.
Materials and Methods
The specimens obtained by Wingate and
Brodkorb in 1960 were collected in asso-
ciation with bones of the extinct crane Grus
latipes, from a vertical fracture on the east-
ern face of the Bermuda Government quar-
ry in Hamilton Parish that was filled with
soil and snail shells of the genus Poecilo-
zonites and was named the “Crane Crev-
ice.”’ The fossil birds, formerly part of the
Pierce Brodkorb collection, are now cata-
loged in the Florida Museum of Natural
History, University of Florida, Gainesville,
and all take the prefix UF PB, which we
have omitted except in the citations of the
357
holotypes and the figure legends. There are
many more specimens available than are
listed among the type material. As para-
types we have listed those specimens that
were used in the descriptions, are illustrat-
ed, or were used for any of the cited mea-
surements. Measurements of long bones of
the limbs usually do not include broken or
juvenile specimens, which accordingly are
not among the paratypes listed. Measure-
ments were taken with digital calipers to
0.01 mm and rounded to the nearest 0.1
mm.
Comparative material examined:—Skel-
etons (complete unless otherwise indicated)
of the following species in the collections
of the National Museum of Natural History,
Smithsonian Institution (USNM), unless
otherwise noted. Amaurolimnas concolor
613963; Coturnicops noveboracensis
556931; Laterallus albigularis 611563; L.
Jamaicensis 492195, 502495; Pardirallus
maculatus 561272—76; Poliolimnas ciner-
eus 560913; Porzana flaviventer 501640,
561276—78, and trunk skeletons 430043,
430931, 430979, 431339—41; Porzana por-
zana 552914; P. albicollis 562750; P. atra
562788; (PS carolina. 501052, 501671; \P.
pusilla 291704-05; Rallus aquaticus
431545, 553039, 553041, UF 34461; R.
limcola. 4899795, S25915; S259 E17, UF
19598, UF 19769, UF 24324, UF 24322; R.
elegans 499437, 525886, 610780; R. lon-
girostris: S258 16529873, 525879; R: Jon-
girostris X R. elegans 525887.
Systematics
Family Rallidae
Genus Rallus Linnaeus
The very long, slender bill of the follow-
ing species clearly places it in the genus
Rallus in its strict sense (Olson 1973a). No
flightless species of the restricted genus
Rallus have been described previously.
Such characters as are preserved in the fos-
sils that are not obscured by flightless ad-
aptations show no evidence of relationship
to the species of Pardirallus (including Or-
358
tygonax), which also have long bills but are
osteologically quite distinct and not espe-
cially closely related to Rallus (Olson
1973a).
Rallus ibycus, new species
Figs. 1-5
Holotype.—Premaxillary symphysis with
anterior portion of internarial bar (pila su-
pranasalis) UF PB5403. Collected in May
1960 by David B. Wingate.
Type locality.—Bermuda, Hamilton Par-
ish, Government Quarry, Crane Crevice.
Chronology.—Middle Pleistocene, pre-
sumably within Oxygen Isotype Stages 13
to 20, approximately 800 to 450 kya (see
discussion).
Measurements (mm) of holotype.—
Length of premaxillary symphysis, 15.5.
Paratypes.—Premaxillary symphyses
5511, 5521. Pila supranasalis 5526. Crania
5401, 5512, 5518. Sterna 5402, 5430. Cor-
acoids 5404, 5415, 5416, 5441, 5456. Hu-
meri 5405, 5417, 5422, 5425-5426, 5428,
5457, 6063—6072, 6073, 6077, 6079. Ulnae
5406, 5423, 5440, 5458. Carpometacarpi
5407, 5419, 5424, 5439, 5459. Pelvis 5429.
Femora 5409, 5432-5438, 5451, 5460, 5
uncataloged specimens. Tibiotarsi 5410,
5448-5449, 5452-5455, 5461, 5491-5492,
5498-5502, 5503, 5541-5542, 5551-5560.
Tarsometatarsi 5442, 5444-5447, 5462,
5543-5546, 6001—6002, 6028-6031, 6055—
6057.
Measurements (mm) of paratypes.—Ros-
trum: length from nasofrontal hinge esti-
mated from 5526 and 5403 to be between
46.5 and 48.0; length of premaxillary sym-
physis 14.1, 15.5, 16.7. Cranium: length
from naso-frontal hinge 28.0; width at post-
orbital processes 15.5, 15.7; width of inter-
orbital bridge 3.3, 3.6, 3.7. Coracoid: length
15.4, 17.4, 18.2, 18.4, 19.5. Sternum: length
along midline (from anterior sternal notch,
not manubrium) 21.2, 21.3; width across
coracoidal sulci 10.7, 12.1; depth of carina
3.7, 4.4. Carpometacarpus: length 14.0,
L6215.1'6.3,,:16.5,)16.95;proxamal -depthi3 .G;
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
4.1, 4.2, 4.3, 4.3. Ulna: length 22.8, 24.5,
25.8, 26.9. Pelvis: sacrum length 27.3, 30.0;
width across antitrochanters 15.4. See also
Vable *1-
Etymology.—Greek, m. ibykos, literally
of a crane, but here a noun in apposition
referring to Ibycus (i: 528. B.C.),"a lyme
poet whose murder was revealed by cranes
(see Anthon 1869). The name is applied to
the new species from its association with
the much larger fossils of cranes, the dis-
covery of which also revealed the extinc-
tion of the rail.
Diagnosis.—Similar in overall size to
Rallus limicola but bill longer and more
slender, with a longer and more decurved
premaxillary symphysis. Flightless, with
the sternum and pectoral elements reduced.
Hindlimb elements much more robust.
Description.—In addition to the longer,
more decurved bill, this species also seems
to have the orbits reduced so the frontal
area in lateral view has the appearance of
sloping more steeply.
Compared with Rallus limicola, the ster-
num is shorter, wider, lacks a manubrial
spine, and has a deep notch between the
coracoidal sulci. The carina is very low and
thick anteriorly, extending laterally as a
rim, which, with the anterior ridge leading
from the keel, creates a deep depression on
the sides of the carina.
The shaft of the coracoid is narrowed,
forming a sharp ridge from the head to mid-
shaft. The procoracoid process is smaller,
more delicate and pointed. The sternocora-
coidal process is much more pronounced
and pointed due to the deep circular inci-
sion in the external margin.
The humerus has the head lower and
smaller, the capital groove deeper, the distal
end narrower, with the tricipital grooves
deeper. The ulna is shorter and stouter and
slightly less curved. The carpometacarpus
is very reduced, short, and stout.
The pelvis in dorsal view is decidedly
wider, both anteriorly and posteriorly, al-
though the ischial area in lateral view is not
as deep as in R. limicola.
VOLUME 113, NUMBER 2
359
Table 1.—Measurements (mm) of selected skeletal elements of two new species of flightless rails from the
Middle Pleistocene of Bermuda.
n Range Mean SD
Rallus ibycus, n. sp.
Humerus
Length 18 28.5—36.3 32.3 ee.
Proximal width 18 6.0—7.2 6.6 0.4
Shaft width 18 1.7—2.2 1.9 0.1
Distal width 18 4.0-4.8 4.4 0.2
Femur
Length 16 34.9-41.5 a7 9 2.6
Proximal width 15 5.7-6.9 6.4 0.3
Distal width [5 5.77.1 6.3 0.5
Tibiotarsus
Length from proximal articular surface 28 45.7-55.5 30/5 3.4
Distal width 27 4.7-5.8 32 0.3
Tarsometatarsus
Length 19 27.1—34.3 30:2 2.4
Proximal width 18 5.1-5.9 5.4 Ome)
Distal width 18 5.1-6.1 5.6 0.3
Porzana pierceéi, n. sp.
Humerus
Length 54 19.7—23.2 24 0:7
Tibiotarsus
Length from proximal articular surface 8 33.6—38.1 33.5 1.6
Distal width Gi 3.13.4 fe 0.1
Tarsometatarsus
Length DS 19.9—23.9 22.3 0.9
Proximal width 25 3:1=3:6 3.4 0.1
Distal width 24 3.43.8 3.6 ot
All of the elements of the hindlimb are
extremely robust compared with R. limico-
la, with heavier shafts and more expanded
articulations. The head of the femur is pro-
portionately larger. The tibiotarsus and tar-
sometatarsus are proportionately shorter.
The cnemial crests of the tibiotarsus are
better developed, the fibular crest is longer,
and the distal tendinal opening is larger.
The tarsometatarsus has the trochleae more
splayed, with the inner trochlea less elevat-
ed and retracted than in R. limicola.
Remarks.—The most likely progenitor of
R. ibycus would be the Virginia Rail, Rallus
limicola, a common migratory species in
eastern North America that has been found
as a “frequent but scarce vagrant’’ in Ber-
muda (Amos 1991:121). Rails described as
being similar to R. limicola are known from
the Pliocene and Pleistocene of North
America (Olson 1977). The Eurasian Water
Rail, Rallus aquaticus, is a less likely an-
cestor on geographic grounds, and also be-
cause of its larger size.
Genus Porzana Vieillot
The following new species of flightless
rail from Bermuda is characterized by very
small size and a short “‘crake-like”’ bill that
invites comparison with the New World
species of Porzana, Laterallus, and Cotur-
nicops. Of particular concern is the Neo-
tropical Yellow-breasted Crake, usually
360
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig, Ie
Comparison of skulls (a—b) and sterna (c—e) of Rallus in lateral view. a, Rallus ibycus, new species
(premaxillary symphysis, holotype UF PB5403; pila supranasalis UF PB5526; cranium UF PB 5401); b, e, R.
limicola female USNM 525915; c, d, R. ibycus UF PB5430, UF PB5402. Scale bar = 2 cm.
known as Porzana flaviventer, for which
Ridgway (1920) once erected the monotyp-
ic genus Hapalocrex (type Rallus flaviven-
ter Boddaert). This species has little resem-
blance to the various species of Porzana
with which it has been placed in most cur-
rent literature. Using mostly external char-
acters, Olson (1970) suggested that it be
placed in the genus Poliolimnas with the
Australo-Malayan species P. cinereus. Al-
though a few authors have accepted Olson’s
conclusion (e.g., Short 1975), it has other-
wise either been widely ignored or even
Sharply attacked (Mees 1982).
In an extensive morphological analysis of
the Gruiformes emphasizing the Rallidae,
Livezey (1998) found the relationships
among the ‘‘crakes’’ and supposed allies to
be difficult to resolve and even more diffi-
cult to reconcile with traditional taxonomy.
The preliminary results of an extensive
sampling of mitochondrial DNA sequences
of the Rallidae indicate that the genus Por-
zana aS now generally construed (e.g., del
Hoyo et al. 1996), is an unnatural assem-
blage (Beth Slikas, National Zoological
Park, pers. comm.). Until the taxonomic
difficulties among the crakes can be better
VOLUME 113, NUMBER 2
361
Fig.:2.
Comparison of skulls (a—b) in dorsal view and sterna (c—e) in ventral view of Rallus. a, Rallus ibycus,
new species (premaxillary symphysis, holotype UF PB5403; pila supranasalis UF PB5526; cranium UF PB
5401); b, e, R: limicola female USNM 525915; c, d, R. ibycus UF PB5430, UF PB5402. Scale bar = 2 cm.
resolved, we have chosen to continue with
general usage in including flaviventer in the
genus Porzana.
The fossil species from Bermuda com-
pares as follows with other crakes from
which it might have been derived. In La-
terallus the nostril is shorter and higher, the
premaxillary symphysis is shorter, and the
interorbital bridge is wider. In species of
Porzana except P. flaviventer, the nostril is
longer and the premaxillary symphysis is
shorter (except in P. pusilla, which has a
long symphysis). In Coturnicops the bill is
extremely short and deep, quite unlike the
fossil species. In the relatively short nostril,
long premaxillary symphysis, and narrow
interorbital bridge, the small Bermuda rail
agrees perfectly with Porzana flaviventer.
A striking feature is the proportions of
the hindlimb. When the hindlimb elements
of Porzana flaviventer are compared with
those of Laterallus jamaicensis it is seen
that whereas the tibiotarsi are of nearly
equal length, the femur in P. flaviventer is
much shorter and all the elements are stout-
er. In Coturnicops noveboracensis the tibi-
otarsus is shorter yet the femur is longer
than in P. flaviventer. Compared with other
362 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Comparison of pectoral and wing elements of Rallus ibycus, new species (a—e) with R. limicola (f-
i, female USNM 525915). a, f, right coracoids in ventral view (a, UF PB5415); b, c, g, left humeri in palmar
view (b, UF PB5422; c, UF PB6072); d, h, left ulnae in internal view (d, UF PB5458); e, i, left carpometacarpi
in internal view (e, UF PB5407). Scale bar = 2 cm.
Fig. 4. Comparison of hindlimb elements in anterior view of Rallus ibycus, new species (a—f) with R. limicola
(gi, USNM 525915 female; j-l, USNM 610783 male). a, left femur UF PB5438; b, right femur UF PB5432;
c, right tibiotarsus UF PB5556; d, left tibiotarsus UF PB5500; e, right tarsometatarsus UF PB5462; f, left
tarsometatarsus UF PB5544. Scale bar = 2 cm.
VOLUME 113, NUMBER 2
363
Fig. 5.
Pelves of rails in dorsal view. a, Rallus ibycus, new species (UF PB5429); b, R. limicola (USNM 610783
male); c, Porzana piercei new species (UF PB5490); d, P. flaviventer (USNM 501640). Scale bar = 2 cm.
species of Porzana, the hindlimb elements
of P. flaviventer are more likewise more ro-
bust, with the femur being proportionately
shorter. As with the cranial elements, the
hindlimb in the small Bermuda rail is most
similar to that of P. flaviventer. The labels
that Pierce Brodkorb left with the speci-
mens indicate that he, too, had concluded
that the small Bermuda rail was derived
from P. flaviventer. The diagnosis of the
new species is therefore based on its dif-
ferences from that species.
Porzana piercei, new species
Figs. 5—9
Holotype.—Complete rostrum UF
PB5413. Collected in March 1960 by
Pierce Brodkorb and David B. Wingate.
Type locality.—Bermuda, Hamilton Par-
ish, Government Quarry, Crane Crevice.
Chronology.—Middle Pleistocene, pre-
sumably within Oxygen Isotype Stages 13
to 20, approximately 800 to 450 kya (see
discussion).
Measurements (mm) of holotype.—
Length from nasofrontal hinge to tip, 16.8;
length of premaxillary symphysis, 5.8.
Paratypes.—Rostra 5421, 5536-5540.
Crania 5412, 5527-5528, 5533. Mandible
5481. Sternum 5414. Scapula 5482. Cora-
coids 5464, 5474, 5483. Humeri 5465,
5484, 5618, 5620, 5620 bis, 5621-5624,
5624 bis, 5625-5629, 5631-5633, 5635-—
5640, 5640 bis?, 5641-5643, 5645-5648,
5650-5651, 5653, 5654-5671, 5686, 6195.
Ulnae 5466, 5485. Carpometacarpi 5476,
5487. Pelvis 5490. Femora 5468, 5478,
5488, 1 uncataloged. Tibiotarsi 5469-5471,
5510, 5547-5550. Tarsometatarsi (all rights
except 5590 & 1 uncataloged with it) 3489,
5575-5585, 5587-5590 + 1 uncataloged,
5594, 5606—5607, 5617, 6060.
Measurements (mm) of paratypes.—
Scapula: length 17.4. Coracoid: length 10.5,
hO:7, Biles Ulna: teneth 16:6, 17.5. Carpo-
metacarpus: length 11.0, 11.2. Pelvis: sa-
crum length 18.2, 19.0; width across anti-
trochanters 11.1, 11.6. Femur: length 25.0,
252, 26.2, 26:2; proximal width 3.8,.4.1,
42, 4.2; distal width 3.7. 3.9, 3.9, 4.1. See
also Table 1.
Etymology.—Dedicated to the late Pierce
Brodkorb who was instrumental in collect-
ing rail material on Bermuda and who first
recognized the affinities of this species.
Diagnosis.—Very similar to P. flaviven-
ter but differing in having the skull and bill
somewhat larger and in being flightless,
with the sternum and pectoral apparatus re-
duced.
Description.—Compared with P. flavi-
venter the cranium is wider, the foramen
364
magnum larger, and the rostrum is wider
with a shorter, wider premaxillary symphy-
sis. The single specimen of mandible, a
symphysis with the left dentary, is abraded
at the tip but the symphysis is wider and
deeper and the bone appears larger than in
P. flaviventer.
The single available sternum lacks the
lateral processes but clearly indicates a
flightless species, as the body of the bone
is much shorter, but slightly wider, with a
much reduced carina that is lower and more
posteriorly situated than in P. flaviventer.
The sternocoracoidal processes are narrow-
er and angled less dorsally in anterior view.
The intercoracoidal notch is wider and the
manubrial spine lacking.
The coracoids are reduced with more
slender shafts and more delicate heads. The
only available scapula has the articular end
and anterior portion of the shaft wider, the
acromion less pointed, and the glenoid facet
larger.
The humeri are only slightly smaller than
in P. flaviventer but have a weaker shaft, a
wider capital groove, and a reduced bicip-
ital crest. The ulnae hardly differ except in
the slightly weaker shafts. The carpometa-
carpi differ only in slightly smaller size and
proportionately shorter intermetacarpal
space.
The pelvis is wider in dorsal view and
the postacetabular portion is proportionate-
ly shorter. Apart from being slightly more
robust, it is difficult to detect any differenc-
es in the hindlimb elements from those of
the living species, particularly given the
very limited comparative material available
for the latter.
Remarks.—This species differs very little
morphologically from Porzana flaviventer.
Although clearly flightless, the degree of re-
duction of the wing and pectoral girdle is
relatively slight compared with many other
flightless rails, probably indicating recency
of derivation.
The Yellow-breasted Crake, P. flaviven-
ter, is a very small, delicate rail that typi-
cally inhabits relatively large bodies of wa-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ter with dense, emergent vegetation. Similar
habitats were unlikely to have been present
in Bermuda, so the birds must have been
able to adapt to different ecological condi-
tions, just as other aquatic rails have adapt-
ed to even harsher insular environments on
Ascension, St. Helena, and Laysan Islands
(Olson 1973b). The modern distribution of
P. flaviventer includes the Greater Antilles
and the Neotropical mainland from south-
ern Mexico to Argentina. An extralimital
record from Antigua suggests that the spe-
cies may have occurred in the Lesser An-
tilles at least up until about 4300 years ago
(Pregill et al. 1988:15).
The Yellow-breasted Crake has never
been found historically in Bermuda (Amos
1991). Although it is possible that P. piercei
was derived from populations of that species
in the Greater Antilles, it seems equally
plausible that the distribution of P. flaviven-
ter may have extended into North America
during periods of the Pleistocene, as was
shown for the Neotropical rail Laterallus ex-
ilis (Olson 1974), in which case colonization
of Bermuda may have been directly from the
North American mainland.
Discussion
Of critical importance is determining the
age of the Bermudian avifauna that includ-
ed at least a crane and a duck with reduced
powers of flight and the two species of
flightless rails described here. Of these fos-
sils Wetmore (1960:1) could say no more
than that “‘it is certain that they are old, and
for the present it is my assumption that they
date back to the Pleistocene.’’ Brodkorb (in
Olson 1977:354) thought that they were
‘‘post-Nebraskan, probably Aftonian or
Kansan.’’ We now know, however, that the
periodicity of sea-level and climatic chang-
es associated with glacial cycles in the
Pleistocene was much more complex than
the traditional view of the four Nebraskan
through Wisconsinan glacial periods, with
their intervening interglacials (e.g., Hearty
1998).
VOLUME 113, NUMBER 2
365
Fig. 6.
Comparison of skulls (a—d, lateral view; b—e, dorsal view) and dorsal views of mandibles (c, f) of
Porzana piercei, new species (a—c) with a very small female P. flaviventer USNM 501640 (d—f). a—b, rostrum,
holotype UF PB5413, and cranium UF PB5412; c, UF PB5481. Scale bar = 2 cm.
There are as yet no direct radiometric
dates on the “‘crane fauna”’ of Bermuda but
circumstantial evidence points very strong-
ly towards a long period of lowered sea lev-
els in the Middle Pleistocene. The ‘“‘crane
fauna” certainly did not survive into the
Holocene as there is no mention of any
such birds in the accounts of early settlers
nor are there fossils of these birds in any
late Quaternary deposits on the island. To
have evolved and sustained such a diverse
endemic avifauna, especially including a
bird as large as a crane, which would have
required extensive savanna-like habitat, the
366 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig: 7:
Comparison of elements of the pectoral girdle of Porzana piercei, new species (a—d) with a very
small female P. flaviventer USNM 501640 (e—-h). a—b, sternum UF PB5414 in lateral and ventral views; c, right
scapula in dorsal view; UF PB5482; d, left coracoid in ventral view UF PB5483. Scale bar = 2 cm.
land area of Bermuda would have had to be
at a maximum for a long period, indicating
a glacial period of lowered sea-level of ex-
ceptional duration. Thus, Brodkorb’s sug-
gestion of Aftonian, which was an intergla-
cial period between the Kansan and Ne-
braskan glacial epochs, can be ruled out.
Interglacials on Bermuda were periods of
marine buildup of carbonate sands on the
flooded Bermuda platform, whereas glacial
periods were marked by sea-levels well be-
low the surface of the platform when de-
position of “terra rossa’”’ soils occured
(Hearty & Vacher 1994). Thus, we would
Fig. 8.
Comparison of wing elements of Porzana piercei, new species (a—d) with a very small female P.
flaviventer USNM 501640 (e—g). a—b, left humeri in anconal view UF PB5637 and UF PB5632; c, right ulna
in internal view UF PB5485; d, right carpometacarpus UF PB5487. Scale bar = 2 cm.
VOLUME 113, NUMBER 2
367
Fig.9.
Comparison of hindlimb elements in anterior view of Porzana piercei, new species (a—f) with P.
flaviventer (g—i, USNM 501640 small female; j—l, USNM 561276 unsexed). a, right femur UF PB5478; b, right
femur UF PB5488; c, right tibiotarsus UF PB5470; d, right tibiotarsus UF PB5510; e, right tarsometatarsus UF
PB9229; f, right tarsometatarsus UF PB5590. Scale bar = 2 cm.
expect deposition of fossils associated with
‘‘terra rossa’’ soils in caves and fissure fills
to have taken place mainly during periods
of lowered sea levels. The longest and most
stable period of lowered sea-levels in the
Quaternary history of Bermuda was from
Oxygen Isotope Stage 13 back to about
Stage 21 (Hearty & Vacher 1994:687, table
1), which spanned a period from about 450
kya to about 800 kya. Although sea-levels
fluctuated during this time, as shown by the
various isotope stages, the amplitude of the
fluctuations was greatly reduced. Thus,
even during the highest stands, the sea was
at or below the level of the Bermuda plat-
form through the preponderance of this in-
terval, during which the subaerial portion
of the platform would have been at its max-
imum extent. It was during this time that
deep limestone dissolution (karstification)
and the accumulation of a massive “‘terra
rossa’’ soil, the Castle Harbour Geosol, oc-
curred (Hearty & Vacher 1994).
This span of several hundred thousand
years of maximum island emergence and
relatively stable terrestrial conditions would
probably have been the only time during
Bermuda’s Pleistocene history when the
“crane fauna”’ could have developed. Sub-
sequent events also provide a very con-
vincing explanation for the disappearance
of this fauna, for it is now known that dur-
ing interglacial Stage 11, 380 to 440 kya,
sea levels rose much higher than had pre-
viously been documented (Hearty et al.
1999), reaching a height of over 20 m
above present level. This would have re-
duced the land area of Bermuda by two or-
ders of magnitude, so that only a few small
islets would have remained. In terms of the
terrestrial biota, this event would essentially
have wiped the slate clean, eliminating the
crane, duck, and rails. A new period of col-
onization and adaption during the next gla-
cial period probably resulted in a substan-
tially different avifauna.
Acknowledgments
We would like to thank the manager and
staff of the Bermuda Government quarry
for alerting Wingate to the extraordinary
deposits of avian fossil material that were
exposed by quarrying operations in 1959
and 1960, and for their considerable coop-
eration and assistance during collection of
368
specimens. We would also like to acknowl-
edge the late Pierce Brodkorb for his part
in collecting, numbering, and studying the
fossil rail material from Bermuda. We thank
Marc Frank and David W. Steadman, Flor-
ida Museum of Natural History, Gaines-
ville, for information, arrangements, access
to fossil and modern collections, lending
specimens, and other considerations. For an
understanding and appreciation of aspects
of the geology of Bermuda we are indebted
to Paul J. Hearty, who also provided many
useful comments on the manuscript. Helen
James generated the statistics in Table 1
from our measurements; the photographs
are by John Steiner and Victor E. Krantz,
Smithsonian Photographic Services, and
these were made ready for publication by
Brian Schmidt, Division of Birds, to all of
whom we are especially grateful. This is
Contribution #26, Bermuda Biodiversity
Project (BBP), of the Bermuda Aquarium,
Natural History Museum and Zoo.
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New Guinea (Merauke and Koembe).—TZoolo-
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Olson, S. L. 1973a. A classification of the Rallidae.—
Wilson Bulletin 85:381—416.
. 1973b. Evolution of the rails of the South At-
lantic Islands (Aves: Rallidae).—Smithsonian
Contributions to Zoology 152:1-—53.
. 1974. The Pleistocene rails of North Ameri-
ca.—Condor 76:169-175.
. 1977. A synopsis of the fossil Rallidae. Pp.
339-373 in S. D. Ripley, Rails of the world: A
monograph of the family Rallidae. Boston, Da-
vid R. Godine, 406 pp.
. 1997. Towards a less imperfect understanding
of the systematics and biogeography of the
Clapper and King rail complex (Rallus longi-
rostris and R. elegans). Pp. 93-111 in R. W.
Dickerman, compiler, The era of Allan R. Phil-
lips: A Festschrift. Albuquerque, New Mexico,
Horizon Communications, 246 pp.
Pregill, G. K., D. W. Steadman, S. L. Olson, & E V.
Grady. 1988. Late Holocene fossil vertebrates
from Burma Quarry, Antigua, Lesser Antil-
les.—Smithsonian Contributions to Zoology
463:1-27.
Ridgway, R. 1920. Diagnoses of some new genera of
bird—Smithsonian Miscellaneous Collections
72(4): 1-4.
Short, L. L. 1975. A zoogeographic analysis of the
South American chaco avifauna.—Bulletin of
the American Museum of Natural History 154:
167-352.
Wetmore, A. 1960. Pleistocene birds in Bermuda.—
Smithsonian Miscellaneous Collections 140(2):
1-11, 3 plates.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):369-—385. 2000.
A new species of Anthias (Teleostei: Serranidae: Anthiinae) from the
Galapagos Islands, with keys to Anthias and eastern
Pacific Anthiinae
William D. Anderson, Jr. and Carole C. Baldwin
(WDA) Grice Marine Biological Laboratory, College of Charleston, 205 Fort Johnson,
Charleston, South Carolina 29412-9110, U.S.A.; (CCB) Department of Vertebrate Zoology,
National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560-0159, U.S.A.
Abstract.—Anthias noeli, new species, is described from 17 specimens col-
lected off the Galapagos Islands in the eastern Pacific, keys to the species of
Anthias and to the species of eastern Pacific Anthiinae are provided, and com-
ments are presented on hermaphroditism in the Anthiinae. The specimens of
the new species, described herein, constitute the first authentic record of An-
thias from the Pacific; the genus is otherwise known only from the Atlantic
where it is represented by seven species. The following characters in combi-
nation allow the separation of Anthias noeli from its congeners: soft rays in
the dorsal fin 15 (rarely 16); tubed lateral-line scales 38 to 46; caudal-peduncle
scales 22 to 25; lower jaw naked or with only a few scales posteriorly; gular
region naked; total gillrakers on first gill arch 37 to 41; dorsal, anal, pelvic,
and caudal fins each with some produced soft rays; anal fin angulate posteri-
orly; depressed anal-fin length 32 to 43% SL; longest dorsal-fin spine (fourth
or fifth) 14 to 20% SL; upper caudal-fin lobe (39-60% SL) longer than lower
(38-57% SL); no teeth on tongue.
During recent collecting trips to the Ga-
lapagos Islands, investigators aboard sub-
mersibles from the Harbor Branch Ocean-
ographic Institution (Fort Pierce, Florida)
have captured a number of new species, in-
cluding 17 specimens of the new anthiine
serranid described herein. Anthiines are
small to medium size brightly colored fish-
es that occur worldwide in shallow to mod-
erate depths of tropical, subtropical, and
temperate seas. They are usually associated
with coral reefs or rocky bottoms that pro-
vide refuge from predators. Individuals of
most species feed upon zooplankton near
the bottom and hastily seek shelter when
approached by predators. Anthiines are of-
ten seen in aggregations, with males attend-
ing large harems. As far as is known, they
are protogynous hermaphrodites, with
many species being sexually dichromatic
and some being sexually dimorphic, espe-
cially in fin structure.
There are approximately 185 valid de-
scribed species in the serranid subfamily
Anthiinae, classified variously in at least 25
genera; additionally, there are a number of
other known undescribed species of Anthi-
inae in museum collections, and there are
undoubtedly other species that remain to be
discovered.
Katayama & Amaoka (1986) restricted
Anthias to include only Atlantic forms, re-
moving Indo-Pacific species more appro-
priately regarded as representatives of
Pseudanthias, Franzia, and Mirolabri-
chthys. Although not clearly stated, it ap-
pears that Katayama & Amaoka (1986:217-—
219, 221) considered Anthias to include the
370
following species: anthias, asperilinguis,
helenensis, menezesi, nicholsi, salmopunc-
tatus, tenuis, and woodsi. We modify their
concept of the genus Anthias to exclude sal-
mopunctatus and tenuis, which, as shown
by Baldwin (1990), appear to be sister spe-
cies and warrant placement in a genus dis-
tinct from Anthias. In addition, we include
in Anthias both the new species described
herein and cyprinoides formerly assigned to
Holanthias by Katayama & Amaoka
(1986).
Although we are not aware of any char-
acter that is clearly synapomorphic for the
species of Anthias, all eight are extremely
similar morphologically and appear to form
a natural group. Herein we provide a di-
agnosis and description of the genus that
distinguishes species of Anthias from all
other serranid fishes.
Methods
Institutional abbreviations follow Leviton
et al. (1985), except for HBOM (=Harbor
Branch Oceanographic Museum). The
methods used here are those of Anderson
& Heemstra (1980), as modified by Ander-
son et al. (1990). Some of those methods
are reiterated or clarified below. Tubed lat-
eral-line scales were counted on both sides
of each specimen of the new species; other
scale counts, except those around the caudal
peduncle, were made on either side, de-
pending on condition of the specimen.
Counts of caudal-peduncle scales were of
the number of scales around the narrowest
part of the peduncle. In making counts of
rows of cheek scales and counts of scale
rows and scales above and below the lateral
line, small scales at orbit and at bases of
dorsal and anal fins were excluded. (Scales
excluded from those counts are distinctly
and abruptly smaller than adjacent scales in
the counted series.) Scales above the lateral
line were counted in a ventroposterior di-
rection from origin of dorsal fin to, but ex-
cluding, the lateral-line scale. Scales below
the lateral line were counted in a dorsoan-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
terior direction from origin of anal fin to,
but excluding, the lateral-line scale. Gillrak-
ers (including rudiments, when present) on
the first gill arch were counted on the right
side. We follow Mabee (1988) and use ‘“‘su-
praneural”’ instead of “‘predorsal’’ to refer
to the short series of bones anterior to the
pterygiophores that support the dorsal fin,
and we conform with Johnson & Patterson
(1993: 557) and Patterson & Johnson
(1995) in using the term “‘epineurals”’’ for
the intermuscular bones that conventionally
have been called ‘“‘epipleurals’”’ in perci-
form fishes. Measurements are given as per-
centages of standard length (SL), except in-
ternarial distance which is also presented as
the quotient of the snout length divided by
the distance between the nares. Osteological
data were recorded from radiographs. Stan-
dard methods for wax histology were used
in preparing the gonadal sections for micro-
scopic examination.
Anthias Bloch
Anthias Bloch, 1792 (type species Labrus
anthias Linnaeus, 1758, by absolute tau-
tonomy).
Aylopon Rafinesque, 1810 (type species La-
brus anthias Linnaeus, 1758, by virture
of the fact that a replacement name re-
tains the type of the prior name; Anthias
incorrectly regarded as preoccupied by
Anthia Weber, 1801, a genus of Coleop-
tera).
Diagnosis.—A genus of Anthiinae dis-
tinguishable from all other genera of the
family Serranidae by the following combi-
nation of characters. No tooth plate on sec-
ond epibranchial. Vertebrae 26 (10 precau-
dal + 16 caudal). Formula for configuration
of supraneural bones, anterior neural spines,
and anterior dorsal pterygiophores 0/0/2/
1+1/1/ (using notation of Ahlstrom et al.,
1976), except A. nicholsi rarely with slight-
ly different placement of supraneural bones.
Principal caudal-fin rays 15 (8 + 7);
branched rays 13 (7 + 6). Dorsal-fin rays
X, 13 to 16 (usually X, 14 or 15, most fre-
VOLUME 113, NUMBER 2
quently X, 15). Anal-fin rays III, 6 to 8
(usually III, 7). Pleural ribs on vertebrae 3
through 10. Epineurals associated with first
11 to 13 vertebrae. Scales ctenoid, with
only marginal cteni (i.e., no ctenial bases
present proximal to marginal cteni—see
Hughes 1981; this type of scale called pe-
ripheral ctenoid by Roberts 1993); no
smaller accessory scales (secondary squa-
mation) at bases of body scales. Most of
head, including maxilla scaly. Lateral line
complete, extending to at least base of cau-
dal fin (running parallel to dorsal body con-
tour a few scale rows ventral to dorsal-fin
base, curving rather abruptly ventral to pos-
terior end of dorsal-fin base to run near
midlateral axis of body). No supramaxilla.
Anterior and posterior nares closely set on
each side of snout; posterior border of an-
terior naris produced into a short flap, but
never into a long filament. No fleshy papil-
lae on border of orbit. Posterior margin of
bony opercle with three spinous processes.
Branchiostegal rays seven. Dorsal fin sin-
gle, not deeply notched between spinous
and soft portions. First caudal vertebra
without parapophyses. Preopercle serrate,
but without antrorse spines. Vomer and pal-
atines with teeth; vomerine tooth patch
without a well-developed posterior prolon-
gation.
Description.—Characters included in the
generic diagnosis form a part of the generic
description and are not repeated. Premax-
illae protrusile. Posterodorsal border of
maxilla not covered by elements of circu-
morbital series when mouth closed. Outer
teeth in jaws mostly conical; inner teeth
mostly villiform or cardiform; some en-
larged as canines. Endopterygoids usually
toothless. Tongue usually without teeth (ex-
cept present on tongue in almost all A. me-
nezesi and A. cyprinoides). Pectoral fin ap-
proximately symmetrical, with 16 to 22
rays. Pelvic-fin rays I, 5. Gillrakers well de-
veloped, total on first arch 37 to 48. Lateral-
line scales 31 to 48. Caudal-peduncle scales
16 to 25. No trisegmental pterygiophores
associated with dorsal and anal fins. No
av
spur on posteriormost ventral procurrent ray
(see Johnson 1975); penultimate ventral
procurrent caudal-fin ray not shortened ba-
sally. Parhypural with well-developed hy-
purapophysis. Autogenous hypurals 5—no
hypural fusions. Epurals 3. Uroneurals |
pair (posterior pair absent). Modified scales
(interpelvic process) overlapping pelvic-fin
bases along midventral line.
Those species of Anthias in which the
larvae have been described lack several
characters found in the larvae of some
American anthiines, viz., specialized larval
scales and serrae on supraoccipital crest,
pterotic ridge, articular, frontal ridge, pari-
etal ridge, and fin spines. In addition, the
known larvae of Anthias have a single ser-
rate supraorbital ridge dorsal to each eye,
in contrast with the larvae of at least two
species of American anthiines that have
three serrate supraorbital ridges on each
side (Baldwin 1990).
Key to the Species of Anthias
la. Lateral-line scales 31—34; sum of lat-
eral-line scales plus total number of
gillrakers on first gill arch, in individual
specimens, 71-76; caudal-fin lobes
moderate (length of upper lobe 31—49%
ST). ae tee. Anthias nicholsi Firth, 1933
(western North Atlantic)
1b. Lateral-line scales 36—48; sum of lat-
eral-line scales plus total number of
gillrakers on first gill arch, in individual
specimens, 75-88; caudal-fin lobes
moderate to well produced (length of
upper lobe 32—110% SL)
2a. Longest dorsal-fin spine (usually the
third) 13-30% SL, 19-30% SL in spec-
imens more than ca. 100 mm SL; third
dorsal-fin spine typically with well de-
veloped filament which may be up to
17% SL; lower caudal-fin lobe usually
longer thamMipperctiink Vite os eee.
Anthias anthias (Linnaeus, 1758)
(eastern Atlantic, including the Medi-
terranean and Adriatic seas)
2b. Longest dorsal-fin spine 10—20% SL;
fin membrane usually extending as a
short filament at tip of each dorsal
aae
SD:
4a.
4b.
Dae
2).0);
6a.
6b.
vey
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
spine, but never produced to the extent
seen in A. anthias; upper caudal-fin
lobe usually longer than lower ...... 3
Soft dorsal-fin rays usually 14 (15 in 1
of 17 specimens); midline of gular re-
gion and lower jaw well covered with
scales; pelvic-fin length 27-41% SL
Anthias
1980
woodsi Anderson & Heemstra,
(western North Atlantic)
Soft dorsal-fin rays usually 15 (rarely
16); gular region naked; lower jaw na-
ked or only partly covered with scales;
pelvic-fin length 33-76% SL
Pectoral-fin rays 19—21 (usually 20 or
21); longest dorsal-fin spine 10-13%
SL; upper caudal-fin lobe 32—44% SL;
pelvic-fin length 33-39% SL; known
only from the eastern South Atlantic SS)
Pectoral-fin rays 17—20 (usually 18 or
19); longest dorsal-fin spine 12—20%
SL; upper caudal-fin lobe 39-—>70%
SL; pelvic-fin length 33-76% SL; not
known from the eastern South Atlantic
pas vps argh a Are Ce Rae ats eR ONE eh 2, lees as 6
Caudal-peduncle scales ca. 18; posteri-
or margin of anal fin rounded; upper
caudal-fin lobe 37-44% SL; no teeth on
tonPUesie eRe Teale tens: Anthias
helenensis Katayama & Amaoka, 1986
(eastern South Atlantic north of the Is-
land of St. Helena)
Caudal-peduncle scales 20—24; poste-
rior margin of anal fin angular; upper
caudal-fin lobe 32-37% SL; tongue
usually with one or two small patches
Of ME Ctlayes.c4-42 ete Fae Sede Anthias
cyprinoides Katayama & Amaoka, 1986
(eastern South Atlantic west of the Is-
land of Pagalu)
Total gillrakers on first gill arch 41—48;
tongue usually with teeth, teeth usually
in narrow elongated patch ... Anthias
menezesi Anderson & Heemstra, 1980
(western South Atlantic)
Total gillrakers on first gill arch 37-41;
tongue usually without teeth
Soft dorsal fin and usually soft anal fin
without produced rays; caudal-peduncle
scales 17 or 18; lateral-line scales 36—
41; two of largest individuals examined
(out of the 10 known specimens) with
teeth on tongue
O:) Bi 56) (6) te) 8: (0;)" enue) 460 10) 26) Ke. (ever ie! jelnle
ue Anthias asperilinguis Giinther, 1859
(western North Atlantic)
7b. Two or more soft dorsal-fin rays and
one or more soft anal-fin rays pro-
duced; caudal-peduncle scales 22-25;
lateral-line scales 38—46; no teeth on
tongue Anthias noeli, new species
(Galapagos Islands, eastern Pacific)
4 je ie je Je) <0
Anthias noeli, new species
Rosy Jewelfish
Figs. 1—5; Tables 1, 2
Material examined.—Seventeen speci-
mens, 62.1 to 173 mm SL; all collected off
the Galapagos Islands in the eastern Pacific
Ocean in depths of 184 to 351 m.
Holotype: USNM 353113 (167 mm SL,
d); seamount SE of Isla San Cristobal—
01°06.48'S, 89°06.70'W; 202 m; Johnson-
Sea-Link-I dive no. 3937; Gilmore & San-
tos, 6 Nov 1995.
Paratypes: USNM 351335 (2, 127-150;
®, 6); off NE Usla” Darwin=01-42 ne
92°00’ W; 351 m; Johnson-Sea-Link-II dive
no. 3103; McCosker & Pawson, 18 Jul
1998. CAS 86573 (1, 132; 3); off Isla Dar-
win—01°41.39'’N, 91°58.88'W; 335 m:;
Johnson-Sea-Link-I dive no. 3963; Mc-
Cosker et al., 20 Nov 1995. USNM 351334
(2, 86-110; 3, 6); off Isla Marchena—
00°24’'N, 90°26.3'W; 303 m; Johnson-Sea-
Link-IT dive no. 3109; Baldwin & Mc-
Cosker, 21 Jul 1998. GMBL 95-34 (1, 139;
2°) & HBOM 107:08471 (1, 123; 2); off
Cabo Douglas, Isla Fernandina—00°17.60'S,
91°39.00'W; 299 m; Johnson-Sea-Link-I
dive no. 3956; Robison & Santos, 16 Nov
1995. CAS 201896 (1, 173; 3); off Isla Pla-
zas (=Plaza Sur in Fig. 2)—00°31'24'S,
90°09'0"W; 308 m; Johnson-Sea-Link-II
dive no. 3096; McCosker & Day, 7 Jul
1998. USNM 351333 (2, 132-170; ?, 3); 2
miles E of Isla Plazas (=Plaza Sur in Fig.
2)—00°32.25'S, 90°09.02’W; 308 m; John-
son-Sea-Link-II dive no. 3096; McCosker
& Day, 7 Jul 1998. CAS 86807 (1, 163;
6); seamount SE of Isla San Cristobal—
01°06.03'S, 89°12.20'W; Johnson-Sea-
VOLUME 113, NUMBER 2
373
Table 1.—Data on morphometric characters for Anthias noeli. Standard lengths are in mm; other measure-
ments, in percentages of standard length.
Character n Min Max. Mean Holotype
Standard length 1? 62.1 173 —— 167
Head, length 17 36.8 43.4 39.4 37.6
Snout, length le 6.0 a2 Iso 8.4
Orbit, diameter 17. el 15.1 135 11.4
Postorbital length of head 17 17.0 20.4 18.8 18.4
Upper jaw, length 17 17.0 19.0 17.9 18.2
Maxilla, width Wi 5.9 7.4 6.6 7.0
Bony interorbital, width Li? 7.6 oJ 8.3 8.6
Internarial distance 17 0.7 LZ 1.0 0.9
Body, depth 17 35.0 42.1 38.5 40.4
Body, width 17 |e | He 15.9 17.6
Predorsal length We 31.8 36.6 34.1 55:7
Prepelvic length LZ 35:7 43.7 39.1 38.2
Preanal length 17 59.4 66.3 63.3 64.0
Caudal peduncle, length ile 21.9 26.7 24.6 20:0
Caudal peduncle, depth 17 11.8 13.6 12.6 13.6
Dorsal-fin base, length 7, 54.4 le), 570 61.7
Pectoral fin, length 17 Deo 33.4 31.2 31.4
Pelvic fin, length 18 32.6 =p 42.7 53.7
Anal-fin base, length 17 16.1 18.5 te 18.2
Anal fin, length £5 323 42.9 37.0 41.9
Upper caudal-fin lobe, length ye 38.8 60.4 49.2 46.9
Lower caudal-fin lobe, length |e) 33.2 57.4 46.8 46.3
First dorsal spine, length 14 6.1 8.5 7.0 7.4
Third dorsal spine, length LS 13s 17.8 153 14.9
Fourth dorsal spine, length it 14.3 19:9 16.6 14.7
Tenth dorsal spine, length is) 10:7 12.6 8 Wea Lee
Longest dorsal spine, length 13 14.3 ee) 16.5 L522
Longest dorsal soft ray, length 11 26.1 44.7 36.0 33.3
Pelvic spine, length 15 15.4 19.8 17-3 5 lf |
First anal spine, length 16 Gad 10.4 8.2 ee |
Second anal spine, length 14 12.0 20.4 15.6 14.4
Third anal spine, length 16 12.8 gre 14.7 12.8
Longest anal soft ray, length 15 24.0 31:8 27.4 31.8
Link-I dive no. 3934; McCosker et al., 5
Nov 1995. UF 110990 (2, 68-134; 2, 9);
seamount SE of Isla San Cristobal—
01°06.23'S, 89°06.91'W; 184-215 m; John-
son-Sea-Link-I dive no. 3935; Robison &
Liberatore, 5 Nov 1995. ANSP 177770 (3
specimens, 62.1-106 mm SL; ?, 2, 3); off
Devil’s Crown, Isla Floreana—01°12.50’S,
90°25.56’W; 232 m; Johnson-Sea-Link-I
dive no. 3945; McCosker & Liberatore, 9
Nov 1995.
Diagnosis.—A species of Anthias distin-
guishable from all other species of the ge-
nus by the following combination of char-
acters. Dorsal-fin rays X, 15 or 16 (usually
X, 15). Anal-fin rays II], 6 or 7 (usually
III, 7). Pectoral-fin rays 18 to 20 (most fre-
quently 19). Lateral-line scales 38 to 46
(usually 39—44). Total gillrakers on first
gill arch 37 to 41 (usually 38—40; no ru-
dimentary rakers). Sum of lateral-line
scales plus total number of gillrakers, on
individual specimens, 78 to 85. Caudal-pe-
duncle scales 22 to 25 (usually 23 or 24).
Caudal fin lunate to deeply forked (larger
individuals tending to have more lunate
fins). Dorsal, anal, pelvic, and caudal fins
each with some produced soft rays. Upper
caudal-fin lobe longer than lower. Anal fin
angulated posteriorly. Gular region without
374
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 2.—Frequency distributions of numbers of fin rays, scales, and gillrakers on first gill arch in Anthias
noeli. Counts for the holotype are indicated by asterisks.
Character 15 16 6
Dorsal soft rays Gs ]
Anal soft rays
Pectoral-fin rays,
Left side
Right side
38 39 40 41
Tubed lateral-line scales,
Left side l
Right side l
i)
Nw NY
£
Circum caudal-peduncle scales l
10 1] 12
Gillrakers
Upper limb 4 La
Lower limb
37 38
Total gillrakers
(upper limb + lower limb) 6
scales. Endopterygoids and tongue without
teeth.
Description.—Characters presented in
the generic and species diagnoses and the
generic description form part of the species
description and are not repeated unless nec-
essary for clarification. Data for morpho-
metric characters appear in Table 1; those
for some countable characters in Table 2.
Mouth nearly terminal, lower jaw exceed-
ing upper jaw very slightly. Maxilla falling
short of vertical through posterior border of
orbit. Maxilla usually widened distally,
with a small shelf or rostrally directed hook
on labial border (as in Anatolanthias api-
omycter—see Anderson et al. 1990:926, fig.
2). Internarial distance 6—12, usually 7—10,
times in snout length. Vertical limb of preo-
percle serrate; horizontal limb smooth or
with several small serrae near angle; angle
with a single spine, a serrated spinous pro-
cess, or relatively large serrae. Distal mar-
gins of interopercle and subopercle usually
Mean
oT, 18 19 20 Mean
15.06
i Se 6.94
ik 14* 2 19.06
2. 10 qr 9.12
43 44 45 46 Mean
4 3%* — 1 41.82
4* 1 4 1 42.06
23 24 DS Mean
i idl y) 2355
27 28 29 30 Mean
3) ih 4 1 28.06
39 40 41 Mean
5 4 l 38.88
smooth or nearly so, occasionally with sev-
eral small serrae.
Premaxilla with inner band of very small
teeth and outer series of larger, mostly an-
teriorly directed, conical teeth; near sym-
physis, two to several teeth along medial
margin of inner band enlarged as posteri-
orly directed conical teeth; outer row of
larger conical teeth usually preceded by one
or two enlarged canine teeth. Dentary with
row of slender conical teeth along lateral
edge of jaw; this row including one to three
teeth, usually enlarged into recurved ca-
nines, at a point approximately one third
length of row from its anterior end; band of
very small teeth extending anteriorly from
this row and reaching to near symphysis;
one to several teeth on inner edge of band
near symphysis enlarged and directed pos-
teriorly; one or two enlarged exserted ca-
nines near symphysis.
Branchiostegal rays seven—anterior
three inserting along ventral edge or ven-
VOLUME 113, NUMBER 2
Pip. 1.
Pacific Ocean. Photograph by Donald Hurlbert, National Museum of Natural History, Smithsonian Institution.
tromedial aspect of hyoid arch, posterior
four inserting on lateral surface of arch.
Pseudobranch well developed, with ca. 22
to 33 filaments; number of filaments in-
creasing with increase in SL (No. filaments
= a+ b[SL]; where a = 17.03, b = 0.08,
f= ©7522).
Most of head, including most of dorsum
of snout, interorbital region, and maxilla,
scaly. Lateral aspect of snout, lachrymal,
branchiostegals, branchiostegal membranes,
and gular region naked (one specimen with
a few scales on lachrymals). Dentary usu-
ally naked, but some specimens with a few
scales posteriorly. Rows of scales on cheek
8 to 11 (usually 9 or 10; difficult to count).
Dorsal and anal fins mostly scaleless, ex-
cept columns of very small scales frequent-
ly present proximally on interradial mem-
branes between soft rays, particularly on
larger specimens. Rows of scales between
lateral line and mid-base of spinous dorsal
fin 2 or 3. Scales between origin of dorsal
Holotype of Anthias noeli, new species, USNM 353113, 167 mm SL; Galapagos Islands, eastern
fin and lateral line 6 to 8. Scales between
origin of anal fin and lateral line 17 to 20.
Longest dorsal-fin spine fourth or fifth.
Second anal-fin spine more robust than first
or third. Procurrent caudal-fin rays 9 (rarely
10) dorsally, 8 or 9 ventrally. Dorsal-fin
membrane produced into a short filament
posterior to each spine. Two or more soft
dorsal-fin rays, one or more soft anal-fin
rays, first two (and on occasion third) soft
pelvic-fin rays (second longest), and cau-
dal-fin lobes produced. Pectoral fin reach-
ing posteriorly to at least as far as vertical
through base of third anal spine, frequently
as far as vertical through base of second
anal soft ray or beyond; dorsalmost pecto-
ral-fin ray always unbranched, ray next to
dorsalmost ray and ventralmost ray occa-
sionally unbranched. Pelvic fin reaching
posteriorly at least to base of first anal soft
ray to as far as well beyond posterior end
of anal-fin base. Pelvic fin inserted beneath
anterior part of pectoral-fin base.
376
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Santa Fe
San Cristobal
]
& _ Champion
Floreano/ . Gardner
Gardner
Espanola
Figs 2.
species. Modified from Grant 1984.
Coloration.—Description based on color
photographs, taken shortly after capture, of
two paratypes (CAS 201896, 173 mm SL;
USNM 351333, 170 mm SL; see Fiesta);
and color notes, made by the second author
in the field, of two other paratypes (USNM
351335, 127 & 150 mm SL). Head mostly
rosy, a yellow streak extending across lach-
Map of Galapagos Islands, eastern Pacific Ocean, showing localities of capture of Anthias noeli, new
rymal and part of cheek and a broad yellow
stripe extending from posterior margin of
eye to posterior tip of opercle. Jaws rosy
except small patch of yellow on upper lip
near premaxillary symphysis. Iris yellow.
Body mostly rosy dorsally, paler ventrally,
a few yellow stripes or blotches on lateral
and ventral aspects of body; black blotch
VOLUME 113, NUMBER 2
present at anterior base of spinous dorsal
fin. Membrane covering dorsal-fin spines
yellow except rosy distally; interradial
membranes mostly rosy; soft dorsal fin
mostly rosy, except membranes separating
three anteriormost dorsal soft rays mostly
yellow or yellow distally and last four rays
pale purple. Anal-fin spines and interradial
membranes yellow; anterior soft anal-fin
rays and interradial membranes yellow ba-
sally, yellow and rosy distally; posterior
portion of soft anal fin rosy to pale purple.
Pectoral fin rosy. Pelvic fin mostly pale
rose, some yellow basally and on mem-
brane between first and second rays. Caudal
fin mostly rosy, some yellow on outer rays
of dorsal and ventral lobes; produced distal
ends of caudal-fin lobes rosy or lilac to pur-
plish in color.
Coloration in alcohol.—Dark spot ante-
rior to base of spinous dorsal fin usually
persisting; dorsum of head frequently
3T7
dusky; fins mostly pallid; remainder of fish
straw-colored.
Sexuality.—Histological examination of
the gonads of all specimens of the new spe-
cies (except for two for which no gonadal
tissue could be found) showed that six in-
dividuals (68.0—-139 mm SL) are females
and nine (86.0—173 mm SL), including the
five largest (150-173 mm SL), are males,
suggesting that Anthias noeli is protogyn-
ous (see section on hermaphroditism in an-
thiines).
Sexual dimorphism.—In specimens more
than about 120 mm SL, females (4 speci-
mens, 123—139 mm SL) have shorter pelvic
fins (33-36% SL vs. 43-55% SL), shorter
longest dorsal soft rays (31-33% SL vs.
38-45% SL), shorter longest anal soft rays
(24-28% SL vs. 28-32% SL), and shorter
depressed anal-fin lengths (33-37% SL vs.
37—42% SL) than do males (5 specimens,
150-173 mm SL).
Fig: 3:
Pacific Ocean.
Paratype of Anthias noeli, new species, USNM 351333, 170 mm SL; Galapagos Islands, eastern
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4.
with only marginal cteni, as in species of Anthias; B,
marginal cteni.
Comparisons.—Anthias noeli is separa-
ble from other species of Anthias as fol-
lows: longest dorsal-fin spine fourth or fifth,
third dorsal-fin spine with at most a short
filament, and upper caudal-fin lobe longer
than lower caudal-fin lobe in A. noeli versus
longest dorsal-fin spine usually third, third
dorsal-fin spine typically with well devel-
oped filament (up to 17% SL), and lower
caudal-fin lobe usually longer than upper
caudal-fin lobe in A. anthias; two or more
soft dorsal-fin rays and one or more soft
anal-fin rays produced and 22 to 25 caudal-
peduncle scales in A. noeli versus soft dor-
sal fin and usually soft anal fin without pro-
duced rays and 17 or 18 caudal-peduncle
scales in A. asperilinguis; upper caudal-fin
lobe length 39 to 60% SL, 18 to 20 (usually
19) pectoral-fin rays, and tongue without
teeth in A. noeli versus upper caudal-fin
lobe length 32 to 37% SE, 20: or 21 pec-
toral-fin rays, and tongue usually with teeth
in A. cyprinoides; 22 to 25 caudal-peduncle
scales and anal fin angulated posteriorly in
A. noeli versus ca. 18 caudal-peduncle
scales and anal fin rounded posteriorly in A.
helenensis; 37 to 41 total gillrakers on first
gill arch and tongue without teeth in A. noe-
li versus 41 to 48 total gillrakers on first
gill arch and tongue usually with teeth in
A. menezesi; 38 to 46 lateral-line scales in
A. noeli versus 31 to 34 in A. nicholsi; 15
or 16 (usually 15) soft dorsal-fin rays and
gular region and dentary without scales
(some specimens with a few scales on den-
Scales of anthiine fishes (posterior fields of scales towards the top of the page). A, ctenoid scale
ctenoid scale with ctenial bases present proximal to
tary posteriorly) in A. noeli versus 14 or 15
(usually 14) soft dorsal-fin rays and midline
of gular region and dentary well covered
with scales in A. woodsi.
In addition, Anthias noeli is distinguish-
able from the other species of Anthiinae
found in the eastern Pacific by the follow-
ing combination of characters: scales cte-
noid, with only marginal cteni (1.e., no cten-
ial bases present proximal to marginal cteni;
see Fig. 4A); maxilla scaly; dentary naked
or with only a few scales posteriorly; gular
region without scales; vomer with teeth; no
teeth on tongue; anterior naris relatively
close to posterior naris (internarial distance
6—12, usually 7—10, times in snout length);
posterior border of anterior naris produced
into a short flap, but never into a long fil-
ament; preopercle without antrorse spines;
urohyal without anteriorly projecting spine;
dorsal-fin rays X, 15 or 16 (usually X, 15);
anal-fin rays III, 6 or 7 (usually III, 7); lat-
eral-line scales 38 to 46; total gillrakers on
first gill arch 37 to 41; diameter of bony
orbit 11 to 15% SL. The most recent com-
parative treatment of the adults of eastern
Pacific anthiines is that of Fitch (1982),
which covers only four of those species, all
found in the northern hemisphere. To facil-
itate identification of all 13 species of east-
ern Pacific Anthiinae, we present a key to
those taxa (see below).
Distribution.—Anthias noeli is known
only from the Galapagos Islands, eastern
VOLUME 113, NUMBER 2
Pacific Ocean, in depths ranging from 184
(0/351. meters (see Fig. (2):
Etymology.—The specific name, noeli, is
for Noel Archambault, IMAX® cameraman/
stereographer, who lost his life on 26 June
1998 in a tragic plane crash in the Gala-
pagos during one of the expeditions on
which the new species was collected. Noel
was a pioneer of modern 3-D film technol-
ogy. It is appropriate to name in his mem-
ory a new species collected using the mod-
ern submersible technology that is allowing
the exploration of oceanic regions previ-
ously inaccessible to scientific study.
Key to the Eastern Pacific Species of
Anthiinae
la. Scales ctenoid, with only marginal
cteni (i.e., no ctenial bases present
proximal to marginal cteni; Fig. 4A)
co lB gi A i ar ea ara 2
1b. Scales ctenoid with ctenial bases pre-
sent proximal to marginal cteni (Fig.
4B) or in one species (Trachypoma
macracanthus) scales cycloid ..... 1
ameeeasilascaly 220.0. Re es 3
Parasillanaked) 2. cae ee eee 5
3a. Vomer edentate; anterior naris some-
what remote from posterior naris; in-
ternarial distance 2.8—3.1 times in
snout length .. Anatolanthias apiomycter
Anderson, Parin, & Randall, 1990
(Nazca Ridge)
3b. Vomerine dentition well developed;
anterior naris relatively close to pos-
terior naris; internarial distance more
than 5.0 times in snout length
4a. Posterior border of anterior naris pro-
duced into a filament, usually reaching
posterior naris when reflected; ventral
surface of dentary and midline of gu-
lar region with scales; bony orbit di-
ameter 7—11% SL (8-11% SL in spec-
imens <160 mm SL, 7-9% SL in
specimens >165 mm SL); teeth usu-
ay present Of tomeuc .. 7... .......-
Pronotogrammus multifasciatus
Gill, 1863 (southern California and
the Gulf of California to northern Peru
and the Galapagos Islands)
4b. Posterior border of anterior naris with-
© * © © » w
5a:
ab:
6a.
6b.
Ta
7b
out a filament, but produced into a
short flap that almost always fails to
reach posterior naris when reflected;
dentary usually without scales, occa-
sionally with a few scales posteriorly;
gular region without scales; bony orbit
diameter 11-15% SL (12-15% SL in
specimens <140 mm SL, 11-13% SL
in specimens >150 mm SL); no teeth
on tongue ...
(Galapagos Islands)
Lateral-line scales 60—71 (usually 62-—
68); specimens more than about 70 mm
SL with sharp spine projecting anteri-
orly from ventral border of urohyal
379
Anthias noeli, new species
. Hemanthias signifer (Garman, 1899)
(southern California to northern Peru)
Lateral-line scales 36-57 (usually 54
or fewer); urohyal without anteriorly
PLOjeCline Spine ii yt .4% eels. 2S
Lateral-line scales 36—40; total gill-
rakers on first arch 38—43; longest
dorsal-fin spine most frequently the
sixth, never the third; third dorsal-fin
spine with short filament, but filament
never well produced; middle rays of
upper and lower lobes of caudal fin
not longest in fin
we Se je eo) Vay car a Le lelve “a ef “ate
Sle “er elise vey eras el Oe se) mt yer ce
Gilbert, 1890 (mid-Gulf of California
[28°N; Fitch 1982] to Colombia
[6.5°N])
Lateral-line scales 49-57; total gill-
rakers on first arch 31-35; longest
dorsal-fin spine—the third; third dor-
sal-fin spine with well produced fila-
ment (at least in larger individuals);
middle rays of upper and lower lobes
of caudal fin longest in fin
a eps a @ el iste) ee 6 oe ees
(Steindachner, 1875) (Hipolito Bank
[27°N], Baja California Sur, in the Pa-
cific and Cabo Lobos [ca. 30°N], So-
nora, in the Gulf of California to Tru-
jillo [8°S], Peru, and the Galapagos Is-
lands [Fitch 1982; Grove & Laven-
berg 1997])
Spines in dorsal fin XI-XIII; ventral
border of preopercle with strong an-
trorse spines
Spines in dorsal fin X; ventral border
of preopercle without antrorse spines,
except in Hypoplectrodes semicinctum
Sa ee ts) eee ee ee a eee a Se oe ee
Pronotogrammus eos
Hemanthias peruanus
380
8a.
8b.
9a.
Ob.
10a.
10b.
iia:
1 1b.
ia:
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
and occasionally in Plectranthias exsul
eel etegte Sec ieee. tee 10
Scales cycloid; spines in dorsal fin
XII; soft rays in anal fin 6
Trachypoma macracanthus
Gunther, 1859 (widely distributed in
the South Pacific, including Easter Is-
land and the Desventuradas Islands off
the coast of Chile [Randall & Cea
Egana 1984; Pequefo & Lamilla
1996a,b; Rojas & Pequefio 1998a])
Scales ctenoid; dorsal spines XI or
XIII; soft rays in anal fin 8 or 9 ... o
Dorsal-fin rays XI, 17 or 18
dee Acanthistius pictus (Tschudi, 1846)
(coasts of Peru and Chile)
Dorsal-fin rays XIII, 14 or 15 ......
Pattee: Acanthistius fuscus (Regan, 1913)
(Easter Island and Sala y Gémez Is-
land [Rojas & Pequeno 1998a])
Soft rays in dorsal fin 19-22 ..... it
Soft. rays m-dorsal’ fin: 1Sror 16242). 12
Preopercle with 2 or 3 antrorse spines
(one spine usually at angle or on ven-
tral margin near angle, other spine[s]
on ventral margin); total gillrakers on
first gill arch 17—20; lateral-line scales
48-55 (most frequently 48-51) ...
Hypoplectrodes semicinctum
(Valenciennes, 1833) (Juan Fernandez
Islands and San Félix Island [in the
Desventuradas Islands] off the coast
of Chile, and possibly Easter Island
[Anderson & Heemstra 1989; Pe-
queno & Lamilla 1996b; Rojas & Pe-
queno 1998a])
Preopercle serrate, but without an-
trorse spines; total gillrakers on first
gill arch 34—37; lateral-line scales 58—
65, usually 61—64 (counts only from
eastemPacilic specimens) =. 4.
.. Caprodon longimanus (Gunther, 1859)
(widely distributed in the Pacific—in-
cluding Easter Island and the Nazca
Ridge and from the Desventuradas
and Juan Fernandez islands off the
coast of Chile, and ‘‘along the coast
of South America”’ [Pequefo & Lam-
illa 1996a:931, 1996b])
Scales on dorsum of head not extend-
ing anterior to nares (Randall 1996:
117); longest soft dorsal-fin ray ca.
26-35% SL; total gillrakers on first
©. ©) ue) je7ne) lel (07a) 0 ici fe
eye) 6. mei ©) e; ‘ei: (eo; (67 fe
gill arch 26—28; in life, two orange-
red bars on body (one just anterior to
anal fin, the other terminating ventral-
ly posteriok toyanal ini a: 5
Plectranthias parini
(Anderson & Randall, 1991) (Sala y
Gomez Ridge and Easter Island
[Randall 1996])
Scales on dorsum of head extending
anteriorly almost to upper lip, except
for triangular premaxillary groove
(Randall 1996:117); longest soft dor-
sal-fin ray ca. 16—>26% SL; total gill-
rakers on first arch 26—31 (usually 28
or 29); in life, brilliant red oblong area
extending from bases of posterior rays
of soft dorsal fin ventrally to just below
midline and then posteriorly over mid-
dle of caudal peduncle to reach mid-
ventral line near base of caudal fin ..
PASUS Midi, MERCER So OE Plectranthias exsul
Heemstra & Anderson, 1983 (Juan
Fernandez Islands and Nazca Ridge;
also reported from Desventuradas Is-
lands by Rojas & Pequeno 1998a)
oe) "ee se Je (ol is, ie: co [0 jo, Se 0:0) 0
1b:
Plectranthias lamillai
Rojas & Pequeno (1998b) described a
new species of Plectranthias, P. lamillai,
from a single specimen (MNHNC P7055,
139.6 mm SL) collected off Alejandro Sel-
kirk Island in the Juan Fernandez Islands
off the coast of Chile. In the description
provided by Rojas & Pequefio (1998b), we
could find no characters that seem to reli-
ably distinguish it from Plectranthias exsul,
also known from off the Juan Fernandez Is-
lands. As a consequence, the first author
made a detailed examination of the holo-
type of P. lamillai (now 136 mm SL) and
compared it closely with two specimens of
P. exsul (USNM 176577—a paratype, 158
mm SL, and USNM 312927, 122 mm SL)
and found that the holotype agrees well
with the original description of P. exsul
and, with one exception, with the speci-
mens of P. exsul with which it was directly
compared. The interorbital region of the
paratype of P. exsul examined in compari-
son is more flattened than that of the ho-
VOLUME 113, NUMBER 2
lotype of P. lamillai; this we attribute to
ontogenetic differences or perhaps to sexual
dimorphism. In contrast, the 122-mm spec-
imen of P. exsul is essentially identical in
head and body shape to the holotype of P.
lamillai. It should be noted that the drawing
with the original description of P. lamillai
(see Rojas & Pequeno 1998b:fig 2) is not a
particularly good representation of the ho-
lotype, especially of the head. Plectranthias
lamillai and P. exsul have somewhat dif-
ferent patterns of coloration, but we think
that the different patterns displayed are best
interpreted as variations on a common
theme. In view of the preceding and pend-
ing a more complete investigation, we deem
it best to consider P. lamillai Rojas & Pe-
queno, 1998, as a junior synonym of P.
exsul Heemstra & Anderson, 1983.
Comments on Hermaphroditism in
Anthiine Serranids
Protogyny has been reported in species
representing a number of anthiine genera,
including Hypoplectrodes (H. huntii [as El-
lerkeldia huntii] and H. maccullochi; Jones
1980 and Webb & Kingsford 1992, respec-
tively), Hemanthias (H. vivanus and H. per-
uanus;,; Hastings 1981 and Coleman 1983,
respectively), Pronotogrammus (P. martin-
icensis [as Holanthias martinicensis|; Co-
leman 1981), Pseudanthias (P. squamipin-
nis [as Anthias squamipinnis|; Fishelson
1970 and Shapiro in a series of studies on
the behavioral aspects of sex reversal—e.g.,
Shapiro 1986), Sacura (S. margaritacea;
Reinboth 1963), and Anthias (A. anthias;
Reinboth 1964). In addition, Heemstra
(1973) provided evidence for protogyny in
Pseudanthias conspicuus (as Anthias con-
spicuus), and Robins & Starck (1961) stated
that Plectranthias garrupellus is probably
protogynous. Thresher (1984) presented a
summary (current through about 1981) of
the information available on the reproduc-
tive biology of anthiines.
One might assume, from the publications
on anthiines cited above, that all of them
381
are protogynous and monandric. Similarly,
based on a number of studies, groupers
(Serranidae: Epinephelinae), in general,
have been considered to display monandric
protogyny, but Sadovy & Colin (1995:961)
found that sexuality in Epinephelus striatus
(the Nassau grouper) “‘is essentially gono-
choristic, with potential for sex change”
and that “‘the juveniles pass through a bi-
sexual stage of gonadal development,”’ thus
illustrating the importance of not making a
priori assumptions about the reproductive
biology of serranid fishes. In a study of the
Serranidae of the eastern Gulf of Mexico,
Bullock & Smith (1991) presented evidence
for protogyny in the anthiine Hemanthias
leptus, but, based on finding a ripening
male of 86 mm SL (p. 21, fig. 8b) and a
sexually mature male of 61 mm SL (p. 207,
pl. Id), entertained the idea that H. leptus
may be diandric, acknowledging, however,
that additional study is needed to confirm
this.
In the subsection entitled Sexuality (un-
der Anthias noeli, new species), we have
presented evidence suggesting that Anthias
noeli is protogynous. To gather a better ap-
preciation of sexuality in the genus Anthias,
William A. Roumillat, at our request, ex-
amined histological sections of the gonads
of 20 specimens of the western Atlantic A.
nicholsi and found 12 females (52.0—125
mm SL), one individual (73.0 mm SL)
transforming from female to male, and sev-
en secondary males (99.9-134 mm SL).
These data strongly suggest that Anthias ni-
cholsi is protogynous. Anthias noeli dis-
plays patterns of gonadal morphology sim-
ilar to those seen in A. nicholsi (Fig. 5),
lending further support to our contention
that A. noeli is probably protogynous.
Sadovy & Shapiro (1987) gave criteria
for diagnosing various types of hermaph-
roditism in fishes. Features that they iden-
tified as strongly indicating protogyny are:
‘*membrane-lined central cavities in testes;
transitional individuals; atretic bodies in
stages 1, 2, or 3 of oocytic atresia within
testes; sperm sinuses in the gonadal wall;
eS)
(o.2)
N
ees
R%
‘oe
Rig. :
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Gonadal sections of specimens of Anthias stained with hematoxylin and eosin-Y. A, A. noeli, new
species, HBOM 107:08471, 123 mm SL, mature female; B, A. noeli, new species, CAS 201896, 173 mm SL,
secondary male; C, A. noeli new species, CAS 86573, 132 mm SL, secondary male; D, A. nicholsi, GMBL 59-
32, 73.0 mm SL, individual transforming from female to male. Abbreviations: AR = artifact; AO = atretic
oocyte; D = dorsal; M = medial; OL = ovarian lumen; OO = oocyte; SS = sperm sinus; TT = testicular
tissue. Photomicrographs by William A. Roumillat, digital imagery by Karen Swanson, South Carolina Depart-
ment of Natural Resources, Charleston.
and experimental production of transitional
or sex-reversed individuals through manip-
ulation of the social system’? (Sadovy &
Shapiro 1987:150). In our studies we found,
with the assistance of Bill Roumillat, that
Anthias nicholsi fulfills all of those criteria,
except the last one which requires live ma-
terial, and that A. noeli meets all but the
last and the one involving transitional in-
dividuals (sensu Sadovy & Shapiro 1987:
147-148) which were not observed. We
feel that it is reasonable to conclude that
both species are protogynous.
Acknowledgments
We appreciate the cooperation of officials
of the Instituto Nacional de Pesca, the
VOLUME 113, NUMBER 2
Parque Nacional Galapagos, and the Esta-
cidn Cientifica Charles Darwin for allowing
the study and capture of fishes, and we ac-
knowledge the support in the field of Al
Giddings of Images Unlimited, David Clark
of David Clark, Inc., the Discovery Chan-
nel, Mandalay Media Arts, LLC, IMAX
Corporation, and the Smithsonian Institu-
tion. David Catania, R. Grant Gilmore, Jr.,
and John E. McCosker provided specimens
and collection data, and Gilmore and
McCosker allowed us to deposit material of
the new species in several museums. Rob-
erto Meléndez very kindly sent the holo-
type of Plectranthias lamillai on loan. Wil-
liam A. Roumillat examined and interpreted
histological sections of the gonads of two
species of Anthias; John McCosker lent us
color transparencies of the new species;
Yvonne Sadovy called our attention to an
important reference; and Antony S. Harold
and Dorian R. McMillan assisted in a num-
ber of ways. Special thanks are due to John-
son-Sea-Link submersible pilots Don Lib-
eratore and Phil Santos (Harbor Branch
Oceanographic Institution), whose expertise
in maneuvering the submersibles and their
collection gear was instrumental in collect-
ing the specimens of the new species. The
black and white photograph of the holotype
of Anthias noeli (Fig. 1) was made by Don-
ald Hurlbert; photomicrographs of the go-
nadal sections of A. noeli and A. nicholsi
(Fig. 5) were provided by W. A. Roumillat,
who also commented upon the section on
hermaphroditism in anthiines. Karen Swan-
son produced the color plate (Fig. 5) of the
gonadal tissues from photomicrographic
transparencies. Support for the publication
of the color plates (Figs. 3 & 5) was pro-
vided by IMAX Corporation, Mandalay
Media Arts, LLC, and the National Muse-
um of Natural History’s Office of Public
Programs. John McCosker and Sarah
Riseman read the entire manuscript and of-
fered suggestions for its improvement. This
is contribution number 165 of the Grice
Marine Biological Laboratory, College of
Charleston.
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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Description of two new blenniid fish species: Entomacrodus lemuria
from the western Indian Ocean and E. williamsi from the western
Pacific Ocean
Victor G. Springer and Ronald A. Fricke
(VGS) Division of Fishes, MRC-159, National Museum of Natural History,
Washington, D.C. 20560-0159, U.S.A.; (RAF) Staatliches Museum fiir Naturkunde in Stuttgart,
Rosenstein 1, D-70191, Stuttgart 1, Germany
Abstract.—Entomacrodus lemuria is described from specimens from Mad-
agascar, Mauritius, and Renunion, and E. williamsi is described from specimens
from Halmahera (Indonesia), Madang (Papua New Guinea), and Duff Islands
(Santa Cruz Islands, Solomon Islands). New distribution records are provided
for E. sealei, a widely distributed Pacific plate endemic, and E. corneliae, a
Marquesas Islands endemic. All four species are members of the Nigricans
Group of Entomacrodus, which group is distinguished from all other blenniids
in having the lateral thirds of the ventral margin of the upper lip crenulate and
the middle third entire. Within the group, these four species form a subgroup
distinguished by having most of the five preopercular pore positions repre-
sented externally by pairs or multiples of pores, as opposed to rarely having
more than one or two positions with pairs or multiples of pores.
Springer (1967) revised the circumtropi-
cal shorefish genus Entomacrodus. There
have been no new species described in the
genus since that publication. Springer
(1972:13) provided additional information
on the Indo-Pacific species, including men-
tion of a variant color pattern in a single
specimen from Madang Harbor, Papua New
Guinea. He tentatively identified the variant
as Entomacrodus sealei Bryan & Herre,
1903, which is otherwise unknown from the
island of New Guinea. Springer (1982:19)
reported a second specimen of the variant
from Ternate, Indonesia, where typical
specimens of E. sealei are also unknown.
He believed that the distribution of the var-
iant, from two localities near the western
margin of the Pacific plate, was allopatric
to that of typical E. sealei, a widely distrib-
uted Pacific plate endemic (Springer 1982:
fig. 8). Recently, in a single collection, J. T.
Williams and associates obtained five spec-
imens of the putative variant together with
a large number of typical specimens of E.
sealei. The sympatry of the two forms con-
vinces us that two species are involved, and
we formally describe the unnamed one
herein. We have also obtained numerous
specimens recently collected from Mauri-
tius, Reunion, and Madagascar of another
undescribed species that also appears to be
related to E. sealei, and we describe that
species herein.
In recent years there has been much in-
terest in documenting (inventorying) the
fish faunas of many parts of the world. Con-
comitantly, there has been much collecting
and specimens have become available from
many localities previously unsampled.
Since 1967, there has been an extensive ac-
cumulation of Entomacrodus specimens in
museum collections, and the genus is in
need of a new revision. We are unable to
undertake that project in the forseeable fu-
ture, but believe that when there is reason-
able certainty that undescribed species ex-
VOLUME 113, NUMBER 2
Table 1.
387
Frequency distributions for number of preopercular pore positions represented by pairs or multiples
of pores in specimens =25 mm SL of the Nigricans Species Group of Entomacrodus. There are five pore
positions.
Number of positions with pairs or
Species N 0 i.
cadenati 84 84 -
textilis a, 54 l
caudofasciatus 149 145 4
vomerinus 113° ,- 409 3
nigricans 343. 314 826
chiostictus 487 184 164
williamsi 7 - -
lemuria 47 - l
sealei 99 - -
corneliae 14 - -
ist, their descriptions are warranted without
delay.
Although the species of Entomacrodus
have not been analyzed cladistically, some
of the seven species groups Springer (1967:
12—13) recognized in the genus are proba-
bly monophyletic. One of these, the Nigri-
cans Group, was defined on the basis of the
morphology of the ventral margin of the
upper lip: the central third of the margin is
entire and the lateral third of the margin on
each side is crenulate. This morphology
does not occur in any other blennioid and,
thus, serves as a reasonable basis for an hy-
pothesis of monophyly of the group. Based
on this character, the two new species are
both members of the Nigricans Group.
Methods
Including the two new species, there are
ten species in the Nigricans Group. There
are very few characters that distinguish
these species, and primary among them are
a few particulars of the color pattern and
sensory pores. For this reason, our descrip-
tions are brief and limited to little more than
multiples of pores
2 3 4 2 X
: = - - 0.00
: . - - 0.02
- - - - 0.03
] - - - 0.04
e) - - = 0:09
DOO ce mah 2 Tee
it 2 3 = Doe
2; aged Seri Oi, LS oa SeBO
- ii Jo. 4.09
- - “ae gh D.00
the essentials necessary to differentiate the
taxa.
Unless defined herein, methods are those
of Springer (1967). All counts of vertical
fin-ray elements were made from radio-
graphs. When split to the base and sup-
ported by a single pterygiophore, the pos-
teriormost anal-fin ray was counted as |
(Method A of Springer 1967:5).
Preopercular pore positions: There are
five sensory canal foramina on the lateral
surface of the preopercle (Springer 1968:
fig. 7; Smith-Vaniz & Springer 1971:fig.
16). Each of these foramina is represented
by (connected to) one or more pores in the
skin immediately external to the foramen.
We term each such representation a “‘preo-
percular pore position’’ or simply “‘posi-
tion’”’ when in context (Fig. 1). Springer
(1967) recognized six pore positions, but
his ventroanteriormost position does not
exit from the lateral surface of the preoper-
cle and is excluded from our definition. The
pores are small and often difficult to see;
however, the number of these positions with
two or more pores is an important character
388 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Piece
Entomacrodus. External limits of preopercle margin
indicated by dotted line; x below ventralmost preoper-
cular pore position; pore position types shown from
ventralmost to dorsalmost: paired, paired, multiples,
multiples, simple.
Diagrammatic illustration of head pores in
for distinguishing species of the Nigricans
Group (Table 1). In specimens shorter than
24 mm SL the pore positions may consist
only or mostly of single pores in all species
of the Nigricans Group; however, in E. le-
muria, all of the five specimens 18.2—22.7
mm SL, we checked for this character, had
two to five positions with pairs of pores,
and the smallest specimen had four posi-
tions with pairs of pores. To be on the con-
servative side, we restrict our characteriza-
tion of pore positions to specimens 24 mm
SL and longer (Table 1).
Color pattern: Many features of the color
pattern in the Nigricans Group species are
highly variable intraspecfically, others are
reasonably consistent within and among the
species. Some of the variation is the result
of poor preservation, but much variation
exists in well-preserved specimens. Among
the most variable markings are the presence
(or absence) and expression of the dusky
bars on the body sides, and some of the
dusky markings on the head and dorsal and
anal fins. These markings, even in speci-
mens that appear to exhibit full expression
of color pattern, are of little use for diag-
nosing species. The more consistent band-
ing of the caudal fin is also of no use. We
have found that only aspects of the color
pattern of the humeral area dorsal to the
pectoral-fin axil and the upper lip are of im-
portance. For these reasons, we restrict our
color-pattern descriptions to these two fea-
tures. The reader desiring more detail is re-
ferred to our illustrations and those of the
Nigricans Group species in Springer (1967:
pls. 18-27).
Institutional abbreviations are those list-
ed in Eschmeyer (1998:16—22).
Materials
Unless noted otherwise, data reported are
taken from specimens cited in Springer
(1967, 197).
Nigricans Species Group
Diagnosis.—Species of Entomacrodus
with: central third of ventral margin of up-
per lip entire and lateral thirds crenulate;
supraorbital cirri branched; nuchal cirri
simple; posteriormost dorsal-fin spine re-
duced, not visible externally; segmented
dorsal-fin rays 13-17 (rarely 13, strongly
modally 14 or 15 in all but one species:
modally 16 in E. vomerinus from the south-
western Atlantic); segmented anal-fin rays
14-18 (rarely 14 and only in E. nigricans,
from the Caribbean; strongly modally 16 in
all but E. vomerinus, modally 17, and 18
only in E. vomerinus); vertebrae 10+23-—26
(strongly modally 34 in all but E. vomeri-
nus, modally 35). Small species, largest
specimen (always male) ranging from 53—
105 mm SL, attaining more than 70 mm
only in E. lemuria (72.1 mm) and E. vom-
erinus (105 mm).
Species Subgroups and Species
Distributions
The ten species of the Nigricans Group
appear to fall into three subgroups based on
the number of preopercular pore positions
389
VOLUME 113, NUMBER 2
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs 3.
humeral spot); b & c, SMNS 213272, 61.5 and 57.4 mm SL.
with pairs or multiples of pores (Table 1).
The monophyly and inter- and intrarelation-
ships of these subgroups are problematic,
but the distributions of each subgroup and
the species within each subgroup seem to
correlate more strongly with geography
than might be expected by chance and may
hint at the naturalness of the groups.
Subgroup 1 comprises five species, of
which 91.5—100% of the specimens have all
pore positions represented by single pores,
and no specimen has more than two posi-
tions represented by pairs or multiples of
pores. Subgroup 2 comprises only one spe-
cies, in which 37.8% of the specimens have
only simple pore positions, and based on
Entomacrodus lemuria, Reunion: a, SMNS 20827, holotype, 59.8 mm SL (pectoral fin obscures
the mean number of positions with paired
or multiple positions (Table 1) is more sim-
ilar to Subgroup 1 than Subgroup 3. Sub-
group 3 comprises four species, in which
no specimen has only simple pore positions.
Subgroup 1 includes all four of the Atlantic
species and one eastern Indian Ocean-west-
ern-and-central Pacific species: E. cadenati
Springer (tropical eastern Atlantic, except
St. Helena and Ascension), E. textilis (Quoy
& Gaimard) (Ascension and St. Helena), E.
nigricans Gill (Caribbean Sea, Bermuda,
Bahamas), E. vomerinus (Valenciennes)
(southwestern Atlantic), and E. caudofas-
ciatus (Regan), eastern Indian Ocean east
to Henderson Island, Pacific Ocean.
VOLUME 113, NUMBER 2
a
39]
Fig. 4. Entomacrodus lemuria, Mauritius: a, USNM 341905, 66.3 mm SL (note damage to second dorsal
fin); b & c, USNM 339747, 43.9 mm SL, ventral and anterior views of head.
Subgroup 2 comprises only E. chiostictus
(Jordan & Gilbert) (eastern Pacific).
Subgroup 3 (Fig. 2) comprises only Pa-
cific or Indian Ocean species: E. corneliae
(Fowler) (Marquesas Islands endemic), E.
sealei (widely destributed Pacific plate en-
demic, but absent from Marquesas Islands),
and the two new species, E. williamsi
(western margin of Pacific plate) and E. le-
muria (southwestern Indian Ocean).
Within Subgroups 1 and 3, all the species
are allopatric except for one co-occurrence
of E. sealei and E. williamsi near the west-
ernmost limits of E. sealei. Considering all
the species together, the only other occur-
rence of sympatry is that of the widely dis-
tributed E. caudofasciatus with E. sealei.
If the sister species of E. lemuria is E.
sealei, or E. sealei and one or both of the
other two Group 3 species, the distribution
pattern shown by E. lemuria and its sister
taxa is another example of eastern Indian
Ocean-Pacific plate disjunct distributions
reported by Springer & Williams (1990).
Those authors hypothesized that the broad
gap between the two distributions was the
result of extinction and explained the gen-
esis of widely distributed Pacific plate en-
demic species, such as E. sealei.
Entomacrodus lemuria, new species
Figs. 3 & 4
Entomacrodus sp. Fricke, 1999:478, Ré-
union; Mauritius.
Holotype.—SMNS 20827, male, 59.8
mm SL, Réunion, west coast, 250 m N
Boucan-Canot, 5 km WSW St. Paul,
21°01'35"S, 55°13'36"E, intertidal area of
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ign.
holotype, 59.8 mm SL lateral and frontal views of head; b & d, USNM 357792, 37.7 mm SL, frontal view of
head and lateral view.
narrow fringing reef, near black rocks on
shore, R. Fricke and S. Ribes, 18 Dec 1998.
Paratypes.—SMNS 21372, 4 specimens,
43.4—61.4 mm SL, collected with the ho-
lotype; AMS 1.39536-001 (5:40.9-61.5 mm
SL), CAS 209025 (5:25.6—60.0 mm SL),
ROM 71976 (5:29.7-54.7 mm SL), RUSI
60472 (5:42.5-63.3 mm SL), USNM
339747 (57:18.1—67.9 mm SL), all Mauri-
tius, W coast, Albion, off rocks at end of
Victoria Avenue, surge zone, O—5 m, A. C.
Gill, P. C. Heemstra, M. Smale, and D. G.
Entomacrodus williamsi, Duff Islands, Santa Cruz Islands, Solomon Islands: a & c, USNM 356864,
Smith, 15 May 1995, field no. PCH 95-
M23; USNM 341905 (4:23.7-66.3 mm
SL), same data as preceding, except: tide
pools, between 25 April and 17 May 1995,
field no. PCH 95-M9; USNM 357266 (29.5
mm SL), Madagascar, Nosy Be, near An-
dilama Beach Hotel, J. Paxton et al., 7 Nov
1988, Vityaz cruise 17, field no. JP 88-9.
Diagnosis.—Ventral margin of upper lip
crenulate on lateral thirds; entire on central
third; nape cirri present, simple; orbital cirri
branched; 1 to 5, usually 3 to 5, preoper-
VOLUME 113, NUMBER 2
393
Fig. 6. Entomacrodus corneliae, Marquesas Islands,
frontal and ventral views of head.
cular pore positions with pairs or multiples
of pores; subquadrate dark blotch present in
humeral area on body (Figs. 3b, c; 4a); up-
per lip frequently with up to 15 alternating
dark and 15 pale bands of subequal width,
up to 7 of each above central, ventrally en-
tire portion of lip (Fig. 4c, d); prominent,
irregular dark mark on head just posterior
to midpostorbital sensory pores (Figs. 3 &
4a).
Description (* denotes character state of
holotype).—Dorsal fin XIII,14 (n = 4),
Pott. 197> (46), > oF XTVA17 © C1) 2) Anal. fin
IE16* (G3) or 11,17.(7/), dast: ray split to
base* (37) or simple (3, of which all have
17 segmented rays). Vertebrae: precaudal
10+23 (1), 24 (32), or 25* (6); pleural ribs
11*-(30); epineural ribs: 13* (9); 14.(6), 15
(11), 16 (0), 17 (1). Segmented pelvic-fin
rays 4* (40). Pectoral-fin rays 13/14 (2), 14/
14* (37), or 14/15 (1). Dorsal procurrent
caudal-fin rays/ventral procurrent caudal-fin
rays 7/6 (2), 7/7* (10), 8/7 (18), 8/8 (4), or
9/8 (1); segmented caudal-fin rays 13* (27),
Nuku Hiva: AMS I.21773010, a, lateral view; b & c,
of which middle 9 are branched, dorsalmost
and ventralmost 2 are simple.
Orbital cirrus branched (usually only on
medial edge of stoutest cirrus), free cirrus
tips 3-8, number tending to increase with
size, variable bilaterally (holotype has 6/5).
Nape cirri 1* on each side, rarely with | or
2 tiny branches. Nasal cirri palmate, 5 to 10
on each side (holotype with 8/9).
Lateral-line a continuous posteroventral-
ly curving tube extending posteriorly to
vertical from base of 9th to 13th dorsal-fin
spine (10th*), continuing along body mid-
side as series of 1 to 8 (5*) tiny, discon-
nected, bipored tubes extending to vertical
from 11th dorsal-fin spine to 5th segmented
ray (2nd segmented ray*). Preopercular
pore positions (Table 1) with 1 to 5* posi-
tions with pairs or multiples of pores.
Color pattern: Highly variable ranging
from almost lacking distinctive markings to
being well marked. Some indication of the
degree of color-pattern variation is indicat-
ed in Fig. 3, all specimens taken in the same
394
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig57.
and frontal views of head.
collection. Most noticeable and consistent
marking a diagnostic subquadrate to ovoid
dark spot on body just dorsoposterior to
pectoral-fin axil. Other markings may in-
clude: irregular dark spot on sensory pore
series posterior to eye; snout above lip and
sides of head with irregular pattern of dark
and pale markings; underside of head with
up to about 6 dusky chevrons separated by
narrower pale bands; upper lip often with
series of up to 15 dark bands alternating
with up to 15 pale bands (dark lip bands,
when present, 5 or 6 on central part of lip
in area over noncrenulate portion of upper
lip. Body with 4 or 5 pairs of dusky bands
on sides, with offset dorsal portions.
Comparisons.—Entomacrodus lemuria
is the only species of Subgroup 3 that has
as the darkest marking on the body, a sub-
quadrate to ovoid spot in the humeral area.
Entomacrodus sealei occasionally has a
dark slash in the humeral region, but it is
Entomacrodus sealei, Eua, Tonga Islands: USNM 335206, 57.8 mm SL, a, lateral view; b & c ventral
much more slender than the humeral spot
of E. lemuria. The humeral spot is essen-
tially the only difference between E. lemu-
ria and E. williamsi. Among the other two
members of Subgroup 3, E. lemuria also
differs from EF. sealei (Fig. 7) in having
more dark and pale lip bands, including
more in the region above the noncrenulate
portion of the upper lip, and in having the
pale bands only a little less slender than the
dark bands (versus almost consistently 7 or
8, each, dark and pale alternating bands,
midlip dark bands much broader than pale
bands, no more than 4 of each in area above
noncrenulate portion of lip). Entomacrodus
lemuria differs from E. corneliae (Fig. 6)
in having only one prominent irregular dark
spot on the head posterior to midpostorbital
sensory pores (versus 2 such dark marks).
Etymology.—From Lemuria, a hypothet-
ical continent, supposed to have existed in
the Indian Ocean and now represented by
VOLUME 113, NUMBER 2
Madagascar and some adjacent islands;
here used as a noun in apposition.
Entomacrodus williamsi, new species
Fig: 5
Holotype.-—USNM 356864, male, 52.8
mm SL, Solomon Islands, Santa Cruz Is-
lands, Duff Islands, Lakao, NW end at Te-
momoa Pt, 09°47'54”S, 165°05'18”E, small
cove, surge channels in big boulders, in
heavy surge, 0-10 m, 24 Sep 1998, J. T.
Williams et al.
Paratypes: USNM 357792, 4 specimens,
36.5—38.1 mm SL, collected with holotype.
USNM 206400, 37.6 mm SL, Papua New
Guinea, Madang Harbour, S edge of Massas
Island, O—2.4 m, 2 Jun 1970, B. B. Collette,
field no. BBC 1501. USNM 356244, 32.4
mm SL, Indonesia, Moluccas, Ternate, Ter-
nate [city], 0.2—-0.5 m, 1-2 July 1979, H.
Singou, field no. HS-F610.
Description (* denotes character state of
holotype).—Dorsal fin XIII,14 (n = 1) or
Poriet>~ (6). Anal fin 11,15 (1), 11,16* ()
onal.t7.(1), last ray split to base* (5) or
simple (2, of which | has 15 and 1 has 17
segmented rays). Vertebrae: precaudal
10+24* (7); pleural ribs 11* (7); epineural
ribs 14* (5) or 15 (1). Segmented pelvic-
fin rays 4* (7). Pectoral-fin rays 13/14 (1)
or 14/14* (6). Dorsal procurrent caudal-fin
rays/ventral procurrent caudal-fin rays 8/2
(hee, 7 (2), 8/8* (2), or 9/8 (1); segmented
caudal-fin rays 13* (7), of which middle 9
are branched, dorsalmost and ventralmost 2
are simple* (6, one specimen damaged).
Orbital cirrus usually branched (usually
only on medial edge of stoutest cirrus), free
cirrus tips 1—6 (1 or 2 probably abnormal),
variable bilaterally (holotype has 5/6). Nape
cirri 1* on each side (1 specimen with cir-
rus forked on one side). Nasal cirri palmate,
4 to 7 on each side (holotype with 6/7).
Lateral-line a continuous posteroventral-
ly curving tube extending posteriorly to
vertical from base of 9th to 12th dorsal-fin
spine (11th*), continuing along body mid-
side as series of 2 to 6 (*5) tiny, discon-
395
nected, bipored tubes extending to vertical
from 13th dorsal-fin spine to 5th segmented
ray (4th ray*). Preopercular pore positions
(Table 1) with 2 to 5 positions with pairs
or multiples of pores (5*; see Table 1).
Comparisons.—The lack of a dark hu-
meral spot in E. williamsi is essentially the
only difference between it and E. lemuria.
There are very few specimens of E. wil-
liamsi, and none of these exhibits distinc-
tive markings on the body, which may have
been present in life. If lack of pattern on
the body of preserved specimens is typical,
this would constitute another difference
from E. lemuria, and the other two species.
Among the other two species of Subgroup
3, E. williamsi also differs from E. sealei
(Fig. 7) in having more dark and pale lip
bands, including more in the region above
the noncrenulate portion of the upper lip
and in having the pale bands only a litte
less slender than the dark bands (versus al-
most consistently 7 or 8, each, dark and
pale alternating bands, midlip dark bands
much broader than pale bands, no more
than 4 of each in area above noncrenulate
portion of lip), and from E. corneliae (Fig.
6) in having only one prominent irregular
dark spot on the head posterior to midpos-
torbital sensory pores (versus 2 such dark
marks).
Etymology.—Named for our colleague
Jeffrey T. Williams, in recognition of his
outstanding efforts in collecting Indo-Pacif-
ic fishes.
Entomacrodus corneliae (Fowler, 1932)
Fig. 6
Additional material (all of the few known
specimens are from the same locality):
Marquesas Islands, Nuku Hiva: AMS
E217 73010) (9).
Entomacrodus sealei Bryan & Herre,
1903
Fig. 7
New distribution records for E. sealei in-
clude: Loyalty Islands: Lifou Island, SMNS
396
21712 (1 specimen). Vanuatu: Tanna,
USNM 344230 (13); Epi, USNM 356399
(7). Fiji: Rotuma, USNM 283060 (1). Ton-
ga: Eua, USNM 329699 (>30), 335206
(12); Tongatapu, USNM 337428 (1);
Vava’u Group, Hunga, USNM 339314 (8);
Vava’u, USNM 340229 (2). Solomon Is-
lands: Santa Cruz Islands, Duff Islands,
Lakao, USNM 356857 (92); Taumako,
USNM 357098 (12); Russell Islands,
Mbanika (or Banika), NMNZ P.26004 (1).
Tuamotu Archipelago: Rangiroa, BPBM
14007 (8). Pitcairn Islands: Henderson Is-
land: BPBM 17091 (10).
Acknowledgments
For the loan of specimens we thank J. E.
Randall (BPBM), M. McGrouther (AMS),
and C. Roberts (NMNZ). Several USNM
colleagues provided a variety of services:
K. Darrow prepared the distribution map;
K. Murphy provided technical assistance; J.
Clayton, L. Palmer, and S. Raredon handled
loan and accession actions. T. B. Griswold
produced the photographs for Figs. 3-7.
The U.S.N.M. Office of Biodiversity Pro-
grams Grant to V. G. Springer and J. T. Wil-
liams provided funds for Williams’s expe-
dition to the Santa Cruz Islands (1998), dur-
ing which important material was collected.
U.S.N.M. Research Opportunities Fund
erant. no. 1233F57A. to..D.G. Smith
(USNM) supported his participation in an
expedition to Mauritius, during which one
of the new species was collected. A draft
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
of this manuscript was reviewed by J. T.
Williams.
Literature Cited
Bryan, W. A., & A. W. C. T. Herre. 1903. Annotated
list of the Marcus Island fishes.—Occasional
Papers of the Bernice P. Bishop Museum 2 (1):
126-139.
Eschmeyer, W. N. 1998. Introduction. Pp. 16—22 in W.
N. Eschmeyer, ed., Catalog of Fishes. Volume
1. California Academy of Sciences, 958 pp.
Fowler, H. W. 1932. The fishes obtained by the Pinchot
South Seas Expedition of 1929, with descrip-
tions of one new genus and three new spe-
cies.—Proceedings of the United States Nation-
al Museum 80 (2906):1—16.
Fricke, R. 1999. Fishes of the Mascarene Islands (Ré-
union, Mauritius, Rodriguez), an annotated
checklist with descriptions of new species.
Koeltz Scientific Books, K6nigstein, 759 pp.
Smith-Vaniz, W. FE, & V. G. Springer. 1971. Synopsis
of the tribe Salariini, with description of five
new genera and three new species (Pisces:
Blenniidae).—Smithsonian Contribtions to Zo-
ology 73:1—72.
Springer, V. G. 1967. Revision of the circumtropical
shorefish genus Entomacrodus (Blenniidae: Sa-
lariinae).—Proceedings of the United States Na-
tional Museum 122 (3582):1—150, Pls. 1—30.
. 1968. Osteology and classisfication of the
fishes of the family Blenniidae.—Bulletin of the
United States National Museum 284:1-—85, pls.
1-11.
. 1972. Additions to Revisions of the blenniid
fish genera Ecsenius and Entomacrodus, with
descriptions of three new species of Ecsenius.—
Smithsonian Contributions to Zoology 134:1—
13%
. 1982. Pacific plate biogeography, with special
reference to shorefishes.—Smithsonian Contri-
butions to Zoology 367:1—82.
, & J. T. Williams. 1990. Widely distributed
Pacific plate endemics and lowered sea level.—
Bulletin of Marine Science 47(3):631—640.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):397—400. 2000.
Ammodytoides leptus, a new species of sand lance
(Teleostei: Ammodytidae) from Pitcairn Island
Bruce B. Collette and John E. Randall
(BBC) National Marine Fisheries Service Systematics Laboratory,
National Museum of Natural History, Washington, D.C., 20560-0153, U.S.A.;
(JER) Bernice P. B. Bishop Museum, P.O. Box 190, Honolulu, Hawaii 96817-0916, U.S.A.
Abstract.—Ammodytoides leptus is described from 23 specimens from Pit-
cairn Island. It has more lateral-line scales than any other known species of
Ammodytoides or Bleekeria (119-123 vs. 88-118). It is thinner than A. pylei,
A. kimurai, and A. gilli (body depth 8.6—9.5% SL vs. 9.5—-11.7%).
During his expedition to the eastern
South Pacific on board the schooner West-
ward (Randall 1978, 1999), the second au-
thor collected the first sand lance recorded
from the South Seas at Pitcairn Island in
January 1971. The purpose of this paper is
to describe this sand lance and assess its
relationships to other members of the genus
Ammodytoides as defined by Ida et al.
(1994), including two additional species de-
scribed by Ida & Randall (1993) and Rand-
all et al. (1994) and A. gilli redescribed by
Collette & Robertson (2000).
Materials and Methods
Type specimens of the new species have
been deposited in the Bernice P. Bishop
Museum, Honolulu (BPBM), the National
Museum of Natural History, Washington,
D.C. (USNM), the Australian Museum,
Sydney (AMS), the Museum of Compara-
tive Zoology, Cambridge (MCZ), and the
National Science Museum, Tokyo (NSMT).
Institutional abbreviations for sources of
comparative material follow Leviton et al.
(1985). Measurements follow Ida and
Randall (1993) and Randall et al. (1994).
Generic nomenclature follows Ida et al.
(1994). All proportions are presented as
percent of standard length (SL).
Ammodytoides leptus, new species
Fig. 1
Ammodytes sp. Randall 1999:24 (an undes-
cribed species from Pitcairn).
Diagnosis.—A species of Ammodytoides
with dorsal-fin rays 50—53; anal-fin rays
24-25; pectoral-fin rays 16—17; pelvic fins
absent; lateral line incomplete, pored later-
al-line scales 114-118 + 4-6 unpored
scales = 119-123; gill rakers on first arch
(6-7) + (22-25) = 29-32; vertebrae (34-—
36) + (26-27) = 61-63, including hypural
plate.
Description.—Body elongate, depth 8.6—
9.5%, width 5.6—6.3%; head length 23.3-
25.1%; snout length 6.7—7.3%; orbit diam-
eter 4.1—4.5%; fleshy interorbital distance
3.5—3.7%; upper jaw length 7.8—8.6%; least
caudal peduncle depth 4.7—5.1%; caudal
peduncle length 6.6—8.3% SL; predorsal
distance 24.0—28.5%; preanal distance
63.3—66.2%; caudal-fin length 13.6—-14.5%;
caudal fin concavity 6.3—7.1%; pectoral-fin
length 9.0—10.1% (Table 1).
Scales small, thin, and cycloid, arranged
in straight diagonal rows; head naked, no
row of small scales on upper part of oper-
cle, scales extending anteriorly to supratem-
poral lateral-line canal; about 10—12 rows
of predorsal scales; fins naked except cau-
dal fin, which has scales extending about
398
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1.—Proportional measurements of type specimens of Ammodytoides leptus expressed in percent of
standard length.
Holotype
Standard length (mm) 96:2 89.1 86.9
Body depth 8.6 a2 95
Body width 6.2 5.8 6.3
Head length 233 24.8 24.4
Snout length 6.8 6.7 70
Orbit diameter 4.2 4.2 4.1
Interorbital space oy 3] Se,
Upper jaw length 7.8 8.0 i9
Caudal peduncle depth 4.7 4.7 4.7
Caudal peduncle length 8.3 7.4 —
Predorsal length 262 24.0 24.7
Preanal length Gna 65.7 66.2
Caudal fin length 14.4 1326 Shed)
Caudal concavity eal 6.7 6.7
Pectoral fin length 10.1 oe wee)
three-fourths distance to posterior margin;
lateral line high on body, ascending from
gill opening to three scales below origin of
dorsal fin, passing posteriorly parallel to up-
per edge of body, pored scales ending high
on body, about 4—6 scales from caudal-fin
base. Suborbital lateral-line canal interrupt-
ed, with four preorbital and four postorbital
pores (as in A. pylei; Randall et al., 1994:
fig. 2A).
In life, dorsum grayish green, sides sil-
very and ventrum iridescent; fins hyaline,
caudal yellowish. Specimens brown in pre-
servative.
Comparisons.—Ammodytoides leptus
has more lateral-line scales than any other
known species of Ammodytoides or Bleek-
eria (119-123 vs. 88-118). It is thinner
Paratypes
86.4 85.3 o20 81.3 79.8 75.8
973 9:0 8.8 8.7 8.9 9:0
6.1 5.6 5.6 6.0 5.6 Do)
24.1 24.2 24.6 24.6 24.4 2a
TA 6.9 7.0 7.0 7.0 LS
4.1 4.1 4.1 4.2 4.4 4.5
Sy 3.6 351 Se EFS) 5
8.1 TS 8.2 8.2 8.0 8.6
4.9 4.9 4.8 4.8 5:0 Sail
6.9 6.8 6.6 6.8 6.8 69
24.3 28.5 PLES 5 24.7 24.4 24.1
66.2 63:3 65.1 65.7 65.8 65.0
13:9 14.2 14.5 Nes ei 13%5 14.1
6.8 6.6 6.6 6.3 6.8 —
9.6 9.0 9.4 9.6 93 9.8
than A. pylei, A. kimurai, and A. gilli (body
depth 8.6—9.5% vs. 9.5—11.7%, Table 2). It
resembles A. gilli and differs from A. pylei
and A. kimurai in several morphometric
characters, larger eye (4.1-5.1% vs. 2.8—
3.7%), longer upper jaw (7.8—9.5% vs. 6.6—
7.9%), longer predorsal distance (24.0—
28.5% vs. 21.8—24.7%), and longer caudal
fin (12.8-16.0% vs. 10.5—12.9%). There is
no row of small scales on upper part of op-
ercle as in A. pylei (Randall et al., 1994).
Ecology.—The series of ten types was
collected from a school of about 25 indi-
viduals seen over sand with high ripple
marks. Schools were fast swimming and ex-
hibited rapid changes in direction, all fish
in perfect unison. When frightened, as by a
spear shot into the school, a few dove into
Fig. L:
20 Jan 1971. Photo by J. E. Randall.
Ammodytoides leptus. Paratype, USNM 360077, 86.9 mm SL, Pitcairn Island, off Gudgeon Point;
VOLUME 113, NUMBER 2
399
Table 2.—Morphometric comparison of four species of Ammodytoides expressed in percent of standard length.
(Data for A. kimurai from Ida & Randall 1993; for A. pylei from Randall et al. 1993; and for A. gilli from
Collette & Robertson 2000).
A. leptus A. kimurai A. pylei A. gilli
Min Max Min Max i iMin,’ | (Max. | °F Mis - | "Max?
Standard length (mm) 75.8 96.2 99.4 121 93.2 168 51.6 84.5
Body depth 8.6 9.5 10.0 10:3 10.0 1t.7 9.5 i
Body width 3:0 6.3 8.1 8.4 7 ee 9.5 5.9 7.8
Head length Pn e&, 25.1 i aes 23:6 22.0 24.6 25.4 27.8
Snout length 6.7 W 6.7 Vike) 6.4 10 6.9 79
Orbit diameter 4.1 4.5 2.8 Dis 2.9 3.7 4.1 52]
Interorbital space 55 Epi 4.1 4.6 Hs) 4.1 ki. 4.9
Upper jaw length 7.8 8.6 TS 7.9 6.6 13 7.8 9.5
Caudal peduncle depth 4.7 ail 4.0 5.0 4.5 ms 4.8 5.4
Caudal peduncle length 6.6 8.3 8.4 9.8 10.6 12.0 3.6 6.0
Predorsal length 24.0 28.5 de | 24.2 21.8 24.7 2 26.9
Preanal length G3:3 66.2 61.3 65.3 63.8 66.8 62.5 65.3
Caudal fin length 13:6 14.5 i I) Re, 10.5 12.0 12.8 16.0
Caudal concavity 6.3 Wl G1 es 5.3 5.8 6.3 es
Pectoral fin length 9:0 10.1 9.1 oS 8.4 92 9.9 yee
N 9 6 10 10
the sand. A larger school (about 100 indi-
viduals) was observed in 30 m around an
offshore rock in the same general area. Am-
modytoides leptus is eaten by larger pelagic
fishes such as the jack, Carangoides ferdau,
as are other species of Ammodytoides.
Etymology.—Named leptus because it is
thinner than other species of Ammodyto-
ides.
Distribution.—Ammodytoides leptus is
presently known only from Pitcairn Island.
Discussion.—Ammodytoides leptus, as
well as two additional species recently de-
scribed by Ida & Randall (1993) and Rand-
all et al. (1994), fits the definition of the
genus as given by Ida et al. (1994) and dif-
fers from species of Bleekeria in the follow-
ing characters: no teeth in jaws; infraorbital
canal interrupted; lateral line ending high
on caudal peduncle instead of curving
downward and continuing onto caudal fin
base; two predorsal bones present; olfactory
rosettes absent; neural and haemal spines
on four posterior caudal vertebrae expanded
and flattened; dorsal-fin rays 51—53 (three
more than the previously recorded range for
the genus), and anal-fin rays 21-25.
In addition to Ammodytoides leptus,
known species of the genus include the
type-species, A. vagus (McCulloch &
Waite, 1916) from Lord Howe Island and
New South Wales; A. renniei (Smith, 1957)
from South Africa, Seychelles Islands, and
the Chagos Archipelago (Winterbottom et
al. 1989, Winterbottom & Anderson 1999);
A. kimurai Ida & Randall, 1993 from the
Ogasawara Islands; A. pylei Randall et al.,
1994 from the Hawaiian Islands; and A. gil-
li (Bean, 1895) from the eastern tropical Pa-
cific (Collette & Robertson 2000).
Material examined.—23 specimens
(71.4-127 mm SL) from 3 original lots
from Pitcairn Island. Holotype USNM
360076 (1, 96.2), off Gudgeon Harbor, 10.5
m, rotenone; J. E. Randall and D. B. Can-
noy; 20 Jan 1971. Paratypes BPBM 16949
(23° 79°8=89: 1)” MCZ>. 157036 (hb, 981.3),
AMS 1.39856-001 (1, 82.9), NSMT-P
59154 (1, 86.4), USNM 360077 (3, 75.8-
86.9), and USNM 360078 (1, 89.8, cleared
and stained), same data as holotype. BPBM
16441 (1, 83.3), dredge haul 2, 48—54 fms;
16 Oct 1967. Additional material examined
but not designated as types due to their poor
condition: BPBM 1660 (12, 71.4—127), off
W. Harbour, stomach of Carangoides fer-
400
dau, 50 ft, J. E. Randall and S. Christian;
27 Dee. 1970:
Comparative material examined: Am-
modytoides kimurai Ida & Randall, 1993.
Paratype. USNM 324610 (1, 121), Japan,
Ogasawara Islands, off Minami-shima, 15
m;.HeIida .& R..L. Pyles) Jun,19972.
Ammodytoides pylei Randall, Ida, and
Earle, 1994. Paratype. USNM 316514 (1,
137), Hawaiian Islands, Oahu, Kahe Point;
RK. LU: Pyle, A. Y. Suzumoto, J.B. Culp: 19
May 1989.
Ammodytoides renniei (Smith, 1957).
RUSI 8440 (1, 56.4), Seychelles Islands,
3°57'S, 54°32”E; 25 Jul 1978. ROM 41487
(1, 60.7), Chagos Archipelago, Peros Ban-
hos Atoll; Isle -dw—Coin; -5°25°20'S:
71°46'52”E; 6 Feb 1979; R. Winterbottom.
Ammodytoides vagus (McCulloch &
Waite, 1916). Holotype. AMS I-9272 (143
mm SL), Lord Howe Island.
Bleekeria mitsukurii Jordan & Evermann,
1903. USNM 59599 (2, 78.5—151), Japan,
Kochi; H. M. Smith. UW 21253 (2, 91.6—
107), Taiwan.
Bleekeria viridianguilla (Fowler, 1931).
Paratypes. ANSP 53462-5 (4, 116—137),
Hong Kong; 1930; G. A. C. Herklots.
Protammodytes sarisa (Robins &
Bohlke, 1970). Holotype ANSP 113091
(115 mm SL) and paratype ANSP 113092
(99.2), Windward Islands, off east coast of
St. Vincent713717 12"Ny 61-05 13) We A877.
m; Pillsbury sta. 874; 6 Jul 1969.
Acknowledgments
Funds for support of field work on board
the Westward were provided to the second
author by the National Geographic Society
(Grant No. 821). We thank curators and
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
staff of museums that house the compara-
tive material for loans and access to the ma-
terial. Ruth Gibbons X-rayed specimens.
The manuscript was reviewed by Thomas
A. Munroe.
Literature Cited
Collette, B. B., & D. R. Robertson. 2000. Redescrip-
tion of Ammodytoides gilli, the eastern Pacific
sand lance (Teleostei: Ammodytidae).—Revista
de Biologia Tropical, (in press).
Ida, H., & J. E. Randall. 1993. Ammodytoides kimurai,
a new species of sand lance (Ammodytidae)
from the Ogasawara Islands.—Japanese Journal
of Ichthyology 40(2):147-151.
, P. Sirimontaporn, & S. Monkolprasit. 1994.
Comparative morphology of the fishes of the
family Ammodytidae, with a description of two
new genera and two new species.—Zoological
Studies, Tokyo 33(4):251—277.
Leviton,. A..E.,..R...H.-Gibbs, Jnj-E. Heald Ga
Dawson. 1985. Standards in ichthyology and
herpetology. Part I. Standard symbolic codes for
institutional resource collections in herpetology
and ichthyology.—Copeia 1985:805-—832.
Randall, J. E. 1978. Marine biological and archaeolog-
ical expedition to southeast Oceania.—National
Geographic Society Research Reports, 1969
Projects:473—495.
. 1999. Report on fish collections from the Pit-
cairn Islands.—Atoll Research Bulletin No.
461, 36 p.
, H. Ida, & J. L. Earle. 1994. Ammodytoides
pylei, a new species of sand lance (Ammody-
tidae) from the Hawaiian Islands.—Pacific Sci-
ence 48(1):80—89.
Winterbottom, R., and R. C. Anderson. 1999. Fishes
of the Chagos Archipelago.—Ecology of the
Chagos Archipelag, C. R. C. Sheppard & M. R.
D. Seaward, eds., Linnean Society Occasional
Publications 2:101—117.
, A. R. Emery, & E. Holm. 1989. An annotated
checklist of the fishes of the Chagos Archipel-
ago, central Indian Ocean.—Royal Ontario Mu-
seum, Life Sciences Contributions No. 145, 226
Pp-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):401—405. 2000.
Neostrengeria binderi, a new species of pseudothelphusid crab from
the eastern Andes of Colombia (Crustacea: Decapoda: Brachyura)
Martha R. Campos
Universidad Nacional de Colombia, Instituto de Ciencias Naturales, Apartado Aéreo 103698,
Bogota, Colombia, S.A.
Abstract.—A new species of freshwater crab of the genus Neostrengeria
Pretzmann, 1965, N. binderi, is described from Alto de Cunday, Tolima De-
partment, Colombia. The addition of this new species brings to 18 species and
2 subspecies the total number of taxa known in this genus endemic to the
Eastern Andes of Colombia.
The genus Neostrengeria Pretzmann,
1965, comprising 18 species and 2 subspe-
cies of pseudothelphusid crabs, is endemic
to the Eastern Andes of Colombia. On the
west slope, the genus was known as far
south as Cundinamarca, but recent collec-
tions in the Cunday region of eastern Toli-
ma have resulted in the discovery of a new
species, which extends the range of the ge-
nus ca. 50 km further south. Additional ex-
plorations in the region have failed to locate
this species west of the Magdalena river,
confirming the association of the genus
with the Eastern Andes.
The systematics of Neostrengeria were
established by Rodriguez (1982) and have
been recently reviewed by Campos (1992,
1994). The geographical distribution of the
genus has been discussed by Campos &
Rodriguez (1985), and Campos (1992,
1994). The general characteristics of the ge-
nus and a key for the identification of the
species was presented by Campos & Le-
maitre (1998).
The terminology used for the different
processes of the male first gonopods is that
established by Smalley (1964) and Rodri-
guez (1982). The material is deposited in
Museo de Historia Natural, Instituto de
Ciencias Naturales, Universidad Nacional
de Colombia, Bogota (ICN-MHN). The ab-
breviations cb and cl indicate carapace
breadth and carapace length, respectively.
Color nomenclature follows Smithe (1975).
Family Pseudothelphusidae Rathbun, 1893
Tribe Strengerianini Rodriguez, 1982
Genus Neostrengeria Pretzmann, 1965
Neostrengeria binderi, new species
Feo 2
Holotype.—Laguna Los Catorce, Vereda
Alto de Cunday, Tolima Department, Co-
lombia, 470 m alt., 22 May 1998, leg. P.
Binder: 1 male, cl 20.7 mm, cb 37.8 mm
(ICN-MHN-CR 1702).
Paratypes.—Same locality data as holo-
type: 2 females, cl /19:15-17.9 mm, :cb 33.6—
31.4 mm (ICN-MHN-CR 1703).
Type locality.—Laguna Los Catorce,
Vereda Alto de Cunday, Tolima Depart-
ment, Colombia, 470 m alt.
Diagnosis.—Carapace with median
groove deep, reaching upper border of
front. First male gonopod with mesial bor-
der slightly convex with strongly subapical
notch in caudal view; apex compressed ce-
phalo-caudal, expanded mesially into sub-
triangular projection; mesial lobe forming
acute triangle.
Description of holotype.—Carapace (Fig.
1A) with cervical groove nearly straight,
deep, ending some distance from lateral
margin. Anterolateral margin with shallow
402
depression behind external orbital angle,
followed by series of papillae on anterolat-
eral half; posterior half smooth. Postfrontal
lobes small, oval, delimited anteriorly by 2
depressions; median groove deep, reaching
upper border of front. Surface of carapace
in front of postfrontal lobes inclined ante-
riorly, depressed towards midline. Front bi-
lobed, lacking distinct upper border in fron-
tal view; lower margin visible in dorsal
view, strongly sinuous in frontal view, with
tubercles. Orbital margins each with row of
tubercles. Dorsal surface of carapace
smooth, covered by small papillae; regions
distinctly marked. Third maxilliped with
merus having sharp angle on distal half of
external margin; exognath 0.57 length of is-
chium (Fig. 1D). Orifice of efferent bran-
chial channel irregularly ovate (Fig. 1B).
First pereiopods heterochelous, left che-
liped larger than the right. Merus with 3
longitudinal crests as follows: upper one
with rows of tubercles, internal lower one
with row of teeth, and external lower one
with row of tubercles. Carpus with few tu-
bercles on internal crest, and blunt distal
spine. Palms of both chelipeds smooth and
swollen; fingers elongate, each 0.58 the
length of propodi, tips crossing, and surfac-
es with rows of small tubercles; finger of
larger chelae not gaping when closed.
Walking legs (pereiopods 2—5) slender
(Fig. 1A). Dactyli elongated, each 1.5 times
as long as propodi, with papillae and 5 lon-
gitudinal rows of large spines diminishing
in size proximally. Number of spines and
papillae on each dactylus arranged as fol-
lows: 1 anterolateral row and 1 anteroven-
tral row each with 6 spines; 1 external row
with 5 spines and 4 intercalated papillae
and | pair of proximal papillae; 1 postero-
lateral row with 4 spines and 1 posterov-
entral row with 5 spines. First gonopod
wide in caudal view, mesial border (Fig.
2A, 1) slightly convex with strongly sub-
apical notch in caudal view. Accesory lobe
(Fig. 2A, 2), shorter than lateral lobe (Fig.
2A, 3), flat caudally, elongate; lateral lobe
wide and semicircular (Fig. 2A—D). Apex
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
in distal view compressed cephalo-caudally
(Fig. 2E), expanded cephalically into sub-
triangular projection; mesial lobe (Fig. 2E,
4) forming acute triangle; mesocaudal pro-
jection of spermatic channel (Fig. 2E, 5) bi-
fid with acute spinules.
Color.—The holotype preserved in al-
cohol is light brown (near 121C, Mikado
Brown) with pale brown (Verona Brown,
223 B) specks on the dorsal side of the car-
apace. The walking legs and chelipeds are
cinnamon brown (Tawny, 38) dorsally and
ventrally. The ventral surface of the cara-
pace is buffy-brown (Antique Brown, 37).
Etymology.—The species is named in
honor of Dr. Philippe Binder, Colombian
scientist who collected the specimens, and
to recognize his efforts in stimulating a new
generation of scientists at the Universidad
de los Andes, Bogota.
Remarks.—This species is most similar
to Neostrengeria aspera Campos, 1992.
Both can be distinguished by features of the
first gonopod. The mesial border of the first
gonopod (in caudal view) of N. aspera is
slightly convex, similar to N. binderi, but
this latter species has a strong subapical
notch. The elongate accesory lobe of N.
binderi is shorter than the lateral lobe,
whereas in N. aspera this lobe is as long as
the lateral lobe, with the apical portion ir-
regular in form and densely covered with
spinules on the distal half (cf., Campos &
Lemaitre 1998). The lateral lobe is spatu-
late, rounded in its distal portion, and sep-
arated from the accesory lobe by a deep
notch in N. aspera, while in N. binderi the
lateral lobe is semicircular and is almost ad-
pressed to the accesory lobe. In the apex of
the gonopod, N. binderi is most similar and
probably closely related to N. gilberti Cam-
pos, 1992. The differences are basically that
in N. gilberti the apex is oval (not com-
pressed cephalo-caudally) and there is an
expansion with a conspicuous cephalic
spine (cf. Campos 1992), whereas in N. bin-
deri the apex consists of a subtriangular
projection. The mesial lobe in N. binderi
shows an acute subtriangular feature, while
VOLUME 113, NUMBER 2 403
Fig. 1. Neostrengeria binderi, new species, male holotype, ICN-MHN-CR 1702. A, dorsal view of carapace
and pereiopods; B, opening of left efferent branchial channel, external view; C, left chela, external view; D, left
third maxilliped, external view; E, frontal view of carapace.
404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Neostrengeria binderi, new species, male holotype, ICN-MHN-CR 1702. A, left first gonopod, caudal
view; B, same, lateral view; C, same, cephalic view; D, same, mesial view; E, same, apex, distal view. 1, mesial
border; 2, accesory lobe; 3, lateral lobe; 4, mesial lobe; 5, mesocaudal projection of spermatic channel.
VOLUME 113, NUMBER 2
in N. gilberti it is smaller and subcircular
with a papilla. The mesocaudal projection
of the spermatic channel in N. gilberti is
awl-shaped, with one spinule on the inner
side, while in N. binderi it is bifid with
acute spinules.
Acknowledgments
The author is grateful to Dr. R. Lemaitre,
G. Styles and Dr. J. Lynch for their useful
comments on this manuscript. The illustra-
tions were prepared by Juan C. Pinzon.
Literature Cited
Campos, M. R. 1992. New species of fresh-water crabs
of the genus Neostrengeria Pretzmann, 1965
(Crustacea: Decapoda: Pseudothelphusidae)
from Colombia.—Proceedings of the Biological
Society of Washington 105:540—554.
. 1994. Diversidad en Colombia de los cangre-
jos del género Neostrengeria.—Academia Co-
lombiana de Ciencias Exactas Fisicas y Natur-
ales. Col. Jorge Alvarez Lleras No. 5:1—143.
405
, & R. Lemaitre. 1998. A new freshwater crab
of the genus Neostrengeria Pretzmann, 1965,
from Colombia (Crustacea: Decapoda: Brach-
yura: Pseudothelphusidae), with a key to the
species of the genus.—Proceedings of the Bio-
logical Society of Washington 111:899—907.
, & G. Rodriguez. 1985. A new species of
Neostrengeria (Crustacea: Decapoda: Pseudoth-
elphusidae) with notes on geographical distri-
bution of the genus.—Proceedings of the Bio-
logical Society of Washington 98:718—727.
Pretzmann, G. 1965. Vorlaufiger Bericht tiber die Fam-
ilie Pseudothelphusidae.—Anzeiger der Oster-
reichischen Akademie der Wissenschaften
Mathematische Naturwissenschaftliche Klasse
(1),1:1-10.
Rathbun, M. 1893. Descriptions of new species of
American freshwater crabs.—Proceedings of
the United States National Museum 16 (959):
649-661, pl. 73-77.
Rodriguez. Gi 1982... Les\ceabes d-ean,) douce
d’Amérique. Famille des Pseudothelphusi-
dae.—Faune Tropicale 22:1—223.
Smalley, A. 1964. A terminology for the gonopods of
the American river crabs.—Systematic Zoology
13:28-31.
Smithe, E B. 1975. Nuturalist’s color guide. The
American Museum of Natural History.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):406—419. 2000.
Freshwater crabs (Brachyura: Potamoidea: Potamonautidae) from
the rainforests of the Central African Republic, Central Africa
Neil Cumberlidge and Christopher B. Boyko
(NC) Department of Biology, Northern Michigan University, Marquette, Michigan 49855, U.S.A.;
(CBB) Division of Invertebrate Zoology, American Museum of Natural History,
Central Park West @ 79th St., New York, New York 10024, U.S.A., and
Department of Biological Sciences, University of Rhode Island,
Kingston, Rhode Island 02881, U.S.A.
Abstract.—A collection of freshwater crabs of the family Potamonautidae
from the Central African Republic made recently by the Central African Re-
public Expedition of the American Museum of Natural History, New York,
comprised four species in two genera. The collection included two species of
Potamonautes MacLeay, 1838, P. paecilei A. Milne-Edwards, 1886, and P.
ballayi A. Milne-Edwards, 1886 and two species of Sudanonautes Bott, 1955,
S. africanus A. Milne-Edwards, 1869, and S. sangha, new species. Only six
species of freshwater crabs have been previously reported to occur in the Cen-
tral African Republic. The addition of S. africanus and S. sangha brings to
eight the number of species of freshwater crabs reported from that country.
The freshwater crabs reported on here
were collected during a systematic survey of
the freshwater and terrestrial Mollusca of the
Central African Republic made by the
American Museum of Natural History. The
survey aimed to document biodiversity, to
catalogue introduced species, and to identify
intermediate hosts that may serve as vectors
of parasite larvae responsible for disease in
humans and domestic animals. The fresh-
water crab collections were made in the vi-
cinity of the village of Bayanga, Central Af-
rican Republic, which lies on the banks of
the Sangha river in a remote and largely
roadless area close to southern Cameroon,
the Republic Populaire du Congo (formerly
French Congo), and Gabon. This region in-
cludes an area of tropical forest that supports
elements of both the West African and Zaire
river basin faunas, and consequently has an
unusually high biodiversity.
The collection of freshwater crabs ob-
tained included two species of Potamonau-
tes MacLeay, 1838, and two species of Su-
danonautes Bott, 1955. Both of these genera
belong to the exclusively African freshwater
crab family Potamonautidae Bott, 1970. The
two species of Potamonautes are P. ballayi
(A. Milne-Edwards, 1886) and P. paecilei
(A. Milne-Edwards, 1886). Both are little-
known species and their diagnostic charac-
ters are redescribed here. Additionally, a lec-
totype is herein designated for P. paecilei.
One of the species of Sudanonautes is
clearly S. africanus (A. Milne-Edwards,
1869). This is a common and well-known
Species that was recently redescribed by
Cumberlidge (1995a, 1999). The second
species of Sudanonautes in the collection is
superficially similar in some respects to S.
africanus, S. chavanesii (A. Milne-Edwards,
1886), S. faradjensis (Rathbun, 1921), S.
floweri (de Man, 1901), S. granulatus
(Balss, 1929) and S. aubryi (H. Milne Ed-
wards, 1853). However, the new specimen
from the Central African Republic has a
number of important characters that do not
conform to the descriptions of any of these
VOLUME 113, NUMBER 2
species (Cumberlidge 1993, 1994, 1995a,
1995b, 1995c, 1995d; 1999). Although the
specimen is a subadult female, and ideally
an adult male is needed to make a definitive
identification, a preliminary description is
provided here, based on several unique so-
matic characters of the specimen. This new
taxon is the eleventh species of Sudanon-
autes to be described (Cumberlidge 1999).
Characters of the gonopods, male abdomen,
and male chelipeds will be described when
more material (including an adult male) be-
comes available.
Only six species of freshwater crabs have
been previously reported to occur in the
Central African Republic (Bott 1955, Cum-
berlidge 1999). These are: Potamonautes
ballayi (A. Milne-Edwards, 1886), P. pae-
cilei (A. Milne-Edwards, 1886), P. dybowski
(Rathbun, 1904), Sudanonautes faradjensis
(Rathbun, 1921), S. floweri (de Man, 1901)
and S. granulatus (Balss, 1929). The addi-
tion of S. africanus and S. sangha new spe-
cies in the present work brings to eight the
number of species of freshwater crab re-
ported from the Central African Republic.
Materials and Methods
Figures were prepared by capturing an
image with a digital camera and completed
using the programs Adobe Photoshop™ and
Adobe Illustrator® (Harvey 1999). The
specimens are deposited in the American
Museum of Natural History, New York,
U.S.A. (AMNH). Abbreviations: Muséum
national d’Histoire naturelle, Paris, France
(MNHN); Muséum royale d’ Afrique central,
Tervuren, Belgium (MRAC); Biology,
Northern Michigan University, Marquette,
Michigan, U.S.A. (NMU); Senckenberg Mu-
seum, Frankfurt, Germany (SMF); cw, dis-
tance across the carapace at the widest point;
cl, carapace length measured along the me-
dian line, from the anterior to the posterior
margin; ch, carapace height (the maximum
height of the cephalothorax); fw, front width
measured along the anterior margin; s, tho-
racic sternite; e, thoracic episternite; s4/s5,
407
S4/s5, s5/s6, s6/s7, s7/s8, sternal sulci be-
tween adjacent thoracic sternites; s4/e4, s5/
e5, s6/e6, s7/e7, episternal sulci between ad-
jacent thoracic sternites and episternites; P1—
P5, pereiopods 1-5.
Systematic Account
Genus Potamonautes MacLeay, 1838
Diagnosis.—Postfrontal crest completely
crossing carapace and meeting anterolateral
margins at epibranchial teeth. Anterolateral
margin always lacking intermediate tooth
between exorbital angle and epibranchial
tooth. Mandibular palp always two-seg-
mented. Exopod of third maxilliped always
with long flagellum. Terminal article of gon-
opod 1 short, about one-quarter to one-third
as long as subterminal segment of gonopod
1. Terminal article of gonopod 2 a long fla-
gellum about 0.5—0.75 times as long as sub-
terminal segment of gonopod 2.
Remarks.—Bott (1955) revised Potamon-
autes and included 38 species and 14 sub-
species, and erected 15 subgenera to accom-
modate these taxa. Since that work, a num-
ber of other species and subspecies have
been described. These are Potamonautes
triangulus (Bott, 1959), P. brincki (Bott,
1960) (Cumberlidge 1994, 1999; Stewart
1997a), P. (Isopotamonautes) anchetiae ma-
chadoi Bott, 1964, P. (Lirrangopotamonau-
tes) lirrangensis adeleae Bott, 1968, P. (J)
senegalensis Bott, 1970, P. dentatus Stew-
art, Coke, & Cook, 1995, P. parvispina
Stewart, 1997b, P. granularis Daniels, Stew-
art, & Gibbons, 1998, and P. reidi Cumber-
hdge; 1999:
Bott (1955) assigned Potamonautes bal-
layi and P. paecilei to the subgenus Longi-
potamonautes Bott, 1955, which also includ-
ed a number of other species of rainforest
river crabs from Central Africa in which
adult males have an elongated, highly arched
right cheliped and sharp teeth on the antero-
lateral margins of the carapace: P. vanden-
brandeni (Balss, 1936), P. schubotzi (Balss,
1914), P. punctatus Bott, 1955, P. ballayi
acristatus Bott, 1955, and P. ballayi gono-
408
cristatus Bott, 1955. Although Bott (1955)
recognised numerous subgenera of Pota-
monautes including Isolapotmonautes, Pla-
typotamonautes, Lirrangopotamonautes and
Longipotamonautes, we prefer here to fol-
low Cumberlidge (1999) and use Potamon-
autes sensu lato for all species, pending a
revision of the entire genus.
Potamonautes ballayi (A. Milne-Edwards,
1886)
Figs. 2
Thelphusa Ballayi A. Milne-Edwards, 1886:
149.—A. Milne-Edwards, 1887:132, pl. 7,
figs. 2., 22)
Potamon (Potamon) ballayi: Rathbun, 1904:
294, pl. 12, fig. 9—Rathbun, 1921:419-
422, pl. 27—28, figs. 1, 10.
Potamon (Potamonautes) ballayi: Balss,
1936:174-177, figs. 9, 12-13.
Potamon ballayi: Chace, 1942:206.—Ca-
part, 1954:827, fig. 3.
Potamonautes (Longipotamonautes) ballayi
ballayi: Bott, 1955:244—245, pl. VII, figs.
2a-d, figs. 23, 73.
Type material and type locality.—Female
holotype, Ngancin (=Nganchu = Ngabé),
Republic Populaire du Congo (formerly
French Congo), 03°18’S, 16°6’E, on oppo-
site bank to Kwamouth, Democratic Repub-
lic of Congo (formerly Zaire), coll. Apr
1884, MNHN.
Material examined.—Central African Re-
public. 1 adult male, cw 18.8 mm (AMNH
17826), about 19 km from the village of
Bayanga, Yobei (Yobé) river, depth 0.1 m,
sandy shore near large dam made entirely of
closely interwoven branches and vines, coll.
J. Cordeiro, 18 Jun 1998.
Diagnosis.—Postfrontal crest not com-
plete, epigastric lobes significantly separated
from postorbital crests, and lateral ends of
postorbital crests not quite meeting antero-
lateral margins (Fig. 1A). Exorbital angle
produced into small pointed tooth; epibran-
chial tooth large, sharp, and pointing for-
ward; anterolateral margin between exorbital
angle tooth and epibranchial tooth smooth,
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
curving slightly outward, lacking interme-
diate tooth; anterolateral margin posterior to
epibranchial tooth smooth, raised, curving
inward over mesobranchial surface of cara-
pace, and not continuous with posterolateral
margin. Carapace height approximately
equal to front width (ch/fw 1.07). Front very
broad, measuring almost one half width of
carapace (fw/cw 0.45) (Fig. 1B). Sidewall of
carapace with distinct vertical sulcus, con-
tinuing downward in pterygostomial region,
dividing sidewall into four parts. Exopod of
third maxilliped with a long flagellum, is-
chium of third maxilliped smooth lacking
vertical sulcus (Fig. 1D). First thoracic ster-
nal sulcus sl/s2 absent; second sulcus s2/s3
deep, running horizontally across sternum;
third sternal sulcus s3/s4 absent so that ster-
num in this region completely smooth (Fig.
1C). Thoracic episternal sulci s4/e4, s5/e5,
s6/e6 and s7/e7 smooth, none marked by
visible groove. Major cheliped of adult
males distinct, with widely arched dactylus
and propodus longer than carapace width
(Fig. 1G). First carpal tooth of inner margin
of carpus of cheliped large, slender, pointed;
second carpal tooth pointed, half size of first
tooth. Lateral inferior margin of merus of
cheliped lined by small teeth, medial inferior
margin of merus of cheliped smooth, with
single large pointed distal meral tooth at dis-
tal end (Fig. 1J); superior surface of merus
ridged by rows of short carinae (Fig. 11).
Terminal article of gonopod 1 short (about
one-third as long as subterminal segment),
longitudinal groove visible on dorsal and su-
perior sides (but not on ventral side); entire
terminal article slim, tubular, and directed
outward at 45° angle to vertical, ending in
wide tip forming distinct pointed process on
medial side; lateral and medial folds on ter-
minal article of gonopod 1 approximately
equal (Figs. 2A—C). The adult size range of
P. ballayi is between cw 28-30 mm.
Description.—For a detailed description
and additional illustrations see Rathbun
(1921) and Bott (1955). For a brief descrip-
tion of the type, see Capart (1954, fig. 3).
Remarks.—Rathbun (1921) recorded the
VOLUME 113, NUMBER 2 409
Fig. 1. Potamonautes ballayi (A. Milne-Edwards, 1886). Male, cw 17.3 mm, AMNH 17826. A, carapace
and eyes, dorsal view; B, cephalothorax, carapace and eyes, frontal view; C, anterior sternum; D, left third
maxilliped; E, left mandible; K abdomen; G, right cheliped, frontal view; H, left cheliped, frontal view; I, carpus
and merus of right cheliped, lateral view; J, carpus and merus of right cheliped, mesial view; K, left second
pereiopod, lateral view. Scale = 1.6 mm (EB), 3.3 mm (D, G—J), and 4.4 mm (A-C, E K).
410
Fig. 2.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
VL
mn csc
oS
a, eR
2 Sty SSeS ORO
Potamonautes ballayi (A. Milne-Edwards, 1886). Male, cw 17.3 mm, AMNH 17826. A, left gonopod
1, cephalic view; B, left gonopod 1, caudal view; C, distal portion of left gonopod 1, superior view; D, left
gonopod 2. Scale = 0.80 mm (C), and 1.67 mm (A, B, D).
mandibular palp as three-segmented. This is
an error, for it is clearly two-segmented (Fig.
1E), as is the case for all members of the
genus, and, indeed all potamonautid African
freshwater crabs (Cumberlidge 1999). Be-
cause the type of P. ballayi is a female, the
gonopods were not illustrated in the first de-
Scriptive works on the species. Gonopod 1
of P. ballayi was subsequently illustrated by
Rathbun (1921) who used a male (cw 26
mm, AMNH 3356) from Stanleyville (now
Kisangani, Democratic Republic of Congo)
and by Bott (1955), who used a male (cw
33 mm, MRAC 17413) from Karawa, Uban-
gi, Democratic Republic of Congo. Gono-
pod 1 of the male from the Central African
Republic is shown here in more detail (Figs.
2A-C), and gonopod 2 of P. ballayi is il-
lustrated for the first time (Fig. 2D).
Ecology.—The specimen from the Central
African Republic was caught in shallow wa-
ter (only 0.1 m deep) near a dam made en-
tirely of closely interwoven branches and
vines. Herbert Lang’s field notes (in Rathbun
1921) record that P. ballayi is common in
shallow forest streams around Stanleyville
(now Kisangani). Lang wrote that it is prob-
able that P. ballayi can live out of water and
that it is only dependent on a certain amount
of moisture. When disturbed, crabs were re-
ported to instantly cover themselves with
mud and secure protection beneath any ob-
ject.
Distribution.—Republic Populaire du
Congo (formerly French Congo), Democrat-
ic Republic of Congo, and Gabon. The type
locality of P. ballayi at Ngabé, Republic Po-
pulaire du Congo lies on the banks of the
Zaire river opposite Kwamouth, Democratic
Republic of Congo. The present study
showed that P. ballayi is present in the Yobé
river, a tributary of the Sangha river which
drains into the Zaire river in a broad marshy
area at Mossaka, Republic Populaire du
Congo. For more localities see Rathbun
(1921), Balss (1936), and Bott (1955).
Potamonautes paecilei (A. Milne-Edwards,
1886)
Fig. 3
Thelphusa paecilei A. Milne-Edwards,
1886:149
VOLUME 113, NUMBER 2
Parathelphusa paecilei: A. Milne-Edwards,
1887:143, pl. 7, figs. 1, la; Ortmann,
1897:300.
Potamon (Parathelphusa) paecilei: Rath-
bun, 1905:257, fig. 167.
Potamon paecilei: Chace, 1942:208.—Ca-
part, 1954:841—842, figs. 34, 37.
Potamonautes (Longipotamonautes) paeci-
lei: Bott, 1955:242-243, pl. VI, figs. 2a—
event figs. 21, 71.
Type material and type locality.—Adult
male lectotype, cw 32 mm (MNHN-B263),
Central Africa, Republic Populaire du Con-
go (formerly French Congo), Latéké (=Lék-
éti), Alima river (14°56’E, 1°36’S), coll. M.
de Brazzae.
Material examined.—Central African Re-
public. 1 adult female, cw 18.6 mm,
(AMNH 17827), 17.3 km from the village
of Bayanga (02°45'43"N, 16°14'12”E), Lossi
creek, depth 1—2 m, bottom of the fine sand
and mud, caught in net in swiftly moving
black water, coll. J. Sullivan, 19 Jun 1998.—
1 adult male, cw 20.7 mm (AMNH 18032),
17.3 km from the village of Bayanga
02645/43'"N, 16°14'12’”E),..Lossi creek,
depth 1—2 m, bottom of fine sand and mud,
caught in net in swiftly moving black water,
coll. M. Lawrence, J. Sullivan, and local res-
idents, 30 Jun 1998.
Diagnosis.—Postfrontal crest either com-
plete or almost complete, wherein lateral
ends of postorbital crests not quite meeting
anterolateral margins (Fig. 3A). Exorbital
angle produced into small, pointed, sharp
tooth; epibranchial tooth large, pointed, di-
rected forward; anterolateral margin be-
tween exorbital angle tooth and epibranchial
tooth smooth, curving outward, lacking in-
termediate tooth; anterolateral margin pos-
terior to epibranchial tooth with two sharp,
forward-pointing teeth; margin otherwise
smooth, continuous with posterolateral mar-
gin. Front very broad, measuring almost
one-half width of carapace (fw/cw 0.45)
(Fig. 3A, B). Carapace height approximately
equal to front width (ch/fw 1.07). Exopod of
third maxilliped with long flagellum, ischi-
411
um of third maxilliped smooth, lacking ver-
tical sulcus (Fig. 3D). First thoracic sternal
sulcus sl/s2 absent; second sulcus s2/s3
deep, running horizontally across sternum;
third sternal sulcus s3/s4 absent; sternum in
this region completely smooth. Episternal
sulci s4/e4, s5/e5, s6/e6 and s7/e7 smooth,
none marked by visible groove. Major che-
liped of adult males distinct, with widely
arched dactylus and a propodus longer than
carapace width. First carpal tooth of inner
margin of carpus of cheliped large, slender,
pointed; second carpal tooth half size of first.
Lateral and medial inferior margins of merus
of cheliped lined by small teeth; single large
pointed distal meral tooth at distal end (Fig.
3J); superior surface of merus ridged by
rows of short carinae (Fig. 31). Terminal ar-
ticle of gonopod 1 short (about one-third as
long as subterminal segment), longitudinal
groove visible on superior side, but not on
dorsal and ventral sides; entire terminal ar-
ticle slim, tubular, curved; directed outward
at an approximately 45° angle to vertical;
ending in broadened upcurved tip; lateral
and medial folds on terminal article of gon-
opod 1 approximately equal size. The adult
size range of P. paecilei is between cw 28-—
30 mm.
Description.—See Bott (1955), and Ca-
part (1954).
Remarks.—One of us (NC) has examined
an adult male syntype (cw 32 mm) of P.
paecilei (MNHN-B 263) from the Alima riv-
er, Latéké French Congo collected by M. de
Brazzae. We designate this specimen here as
the lectotype. A different male syntype (now
paralectotype) of P. paecilei (cw 22.7 mm)
was figured by Capart (1954). The charac-
ters of this species (Fig. 3) include a large
forward-pointing epibranchial tooth; a large
tooth (or two teeth) behind the epibranchial
tooth on the anterolateral margin; an en-
larged major cheliped in adult males with a
widely arched dactylus and a propodus that
is longer than the carapace width; a long
sharp distal meral spine on the merus of the
cheliped; and the ischium of the third max-
412 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Potamonautes paecilei (A. Milne-Edwards, 1886). Female, cw 18.6 mm, AMNH 17827. A, carapace
and eyes, dorsal view; B, cephalothorax, carapace and eyes, frontal view; C, anterior sternum; D, left third
maxilliped; E, left mandible; EK abdomen; G, right cheliped, frontal view; H, left cheliped, frontal view; I, carpus
and merus of right cheliped, lateral view; J, carpus and merus of right cheliped, mesial view; K, left second
pereiopod, lateral view. Scale = 2.2 mm (E), 3.3 mm (D, I, J), 4.4 mm (A, B, F—-H, K), and 5.9 (C).
VOLUME 113, NUMBER 2
illiped is smooth and a vertical sulcus is
lacking.
The sidewall of the carapace of the spec-
imens from the Central African Republic has
a distinct vertical sulcus which continues
downward across the pterygostomial region,
dividing the carapace sidewall into four
parts. This contrasts with the type from the
Republic Populaire du Congo where the
sidewall of the carapace is divided into only
three parts.
Ecology.—The specimens from the Cen-
tral African Republic were netted in a small
stream (1-2 m deep), with swiftly moving
black water flowing over fine sand and mud.
Distribution.—Republic Populaire du
Congo, and Democratic Republic of Congo.
The Yobé river is a tributary of the Sangha
river which drains southwestern Central Af-
rican Republic and forms part of the border
between the Central African Republic and
Cameroon, and then between Cameroon and
Republic Populaire du Congo, before joining
the Zaire river in a broad marshy area at
Mossaka, Republic Populaire du Congo. The
type locality lies on the Alima river which
flows into the Zaire river just to the south of
Mossaka and contributes to the same ex-
panse of marsh and wetlands as the Sangha
river.
Sudanonautes Bott, 1955
Sudanonautes Bott, 1955:295.—Cumberlid-
ge, 1999:172-176.
Diagnosis.—Intermediate tooth on antero-
lateral margin between epibranchial tooth
and exorbital angle. Postfrontal crest prom-
inent, almost horizontal, complete, with lat-
eral ends meeting anterolateral margins. Car-
apace sidewall divided by two sulci into
three parts. Mandibular palp two-segmented;
terminal segment consisting of large oval
posterior lobe (in three species there a small
but distinct anterior process at junction be-
tween segments). Long, plumose flagellum
on exopod of third maxilliped in all species.
Sternal sulcus s3/s4 represented only by two
short notches at sides of sternum. Terminal
413
article of gonopod | very long (at least two-
thirds as long as subterminal segment). Ter-
minal article of gonopod | either slim and
needle-like (where longitudinal groove not
visible) or broadened in middle (the result
of a higher medial fold) with longitudinal
groove visible at least for part of length. Ter-
minal article of gonopod 2 very short, one-
fifteenth length of subterminal segment.
Distribution. —The genus is present in
Céte-d’ Ivoire, Ghana, Togo, Benin, Nigeria,
Cameroon, Gabon, Bioko (Fernando Po),
Central African Republic, Congo, Zaire,
northern Angola, and southwest Sudan. The
eleven species of Sudanonautes are found in
the inland waters of West and Central Africa
in a region bounded by Céte-d’ Ivoire, south-
west Sudan, and northern Angola. This area
includes the Upper Guinea rainforests, the
Lower Guinea forest together with the sa-
vannas of the eastern part of West Africa,
and the offshore island of Bioko. In Central
Africa seven species of Sudanonautes (S. af-
ricanus, S. aubryi, S. floweri, S. granulatus,
S. chavanesii, S. faradjensis, and S. sangha)
share the rivers and forests with species of
Potamonautes and Erimetopus A. Milne-Ed-
wards, 1886 (Bott 1955, Cumberlidge 1999).
Sudanonautes africanus (A. Milne-
Edwards, 1869)
Thelphusa africana A. Milne-Edwards,
1869:186, pl. XI, figs. 2, 2a,b.—A. Milne-
Edwards, 1887:124—126, pl. IV, fig. 8.
Potamon (Potamonautes) africanus: de
Man, 1903:41, pl. [X, figs. 7—9.—Rath-
bun, 1904, pl. 16, fig. 6—Rathbun, 1905:
188-190, fig. 47—Balss, 1929:124—125,
figs. 5—7.—Balss, 1936:166.
Potamon (Potamonautes) africanum: Colo-
si, 1920:34.—Colosi, 1924:21, fig. 16.—
Roux, 1927:237.
Potamon africanus: Chace, 1942:204.—Ca-
part, 1954:824, figs. 1, 6.
Sudanonautes (Sudanonautes) africanus af-
ricanus: Bott, 1955:295-—298, figs. 61, 93—
95, 103 a-d, pl. 24, figs. 2a—c, 3.—Bott,
1959:1004—1005.—Monod, 1977:1216
414
(not figs. 93-95, 102).—Monod, 1980:
384, pl. V, fig. 27.
Sudanonautes africanus: Cumberlidge,
1995a:588—598, figs. 1-3, table 1.—Cum-
berlidge, 1999:181—184, figs. 30B, 32B,
33B, 34B, 35C, 36E, 37C, 53P, 54-57,
60B, 67A, table IX.
Type material.—The holotype used by A.
Milne-Edwards (1869) to describe Thelphu-
sa africana was a small juvenile (MNHN)
(cw 17 mm) collected from Gabon by M.
Aubry-Lecomte. A more detailed descrip-
tion by A. Milne-Edwards (1887) was based
on a larger, but still subadult female
(MNHN) (cw 53 mm) collected from the
river Ogoué, Congo (=Gabon). Because nei-
ther of these specimens was suitable to re-
describe the species (one is a juvenile and
the other a subadult female), and because no
topotypes were available, the species was re-
described by Cumberlidge (1995a) from an
adult male (cw 83 mm) from Cross River
State, Nigeria (NMU 9.IV.1983), and an
adult female (cw 108 mm) from a tributary
of the Ikpan river, Cross River State, Nigeria
(NMU 5.IV.1983).
Material examined.—Central African Re-
public. Ten specimens, sub-adults and juve-
niles (no adults) (AMNH 18033), 19.5 km
from the village of Bayanga (03°05'27'N,
16°16’40"E), Mapoyo (Mboyé) creek, depth
1—1.5 m, either on muddy bottom, or in bur-
rows in overhanging banks, coll. M.
Lawrence, J. Sullivan, and local residents, 17
Jun 1998. Four specimens, 1 adult male and
3 juveniles (AMNH 18034), about 5 km from
the village of Bayanga, Mobeya [Moubia?]
creek, upstream of mouth, depth 0.5—1 m,
sandy, gravely and muddy pools, in burrows
in overhanging banks, among roots, under
logs, coll. M. Lawrence and local residents,
26 Jun 1998. One subadult male (AMNH
18035), 19.5 km from the village of Bayanga
(03°05'27"N, 16°16'40"E), Mapoyo (Mboyé)
creek, depth 1—1.5 m, on muddy bottom, or
in burrows in overhanging banks, coll. M.
Lawrence, J. Sullivan, and local residents, 17
Jun 1998. Four specimens, all juveniles
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
(AMNH 18036), about 5 km from the village
of Bayanga, Mobeya [Moubia?] creek, up-
stream of mouth, depth 0.5—-1 m, sandy,
gravely and muddy pools, in burrows in over-
hanging banks, among roots, under logs, coll.
M. Lawrence, J. Sullivan, and J. B. Kindi-
moungo, 28 Jun 1998.
Type locality.—Gabon.
Diagnosis.—Carapace relatively flat (ch/
fw 1.06). Postfrontal crest smooth almost
straight; spanning entire carapace, meeting
anterolateral margins at epibranchial teeth;
posterior surface of carapace in cardiac and
branchial regions with patches of raised cir-
cular blisters, lateral parts with fields of
raised short lines (carinae); semi-circular,
cardiac, urogastric grooves very deep. Prox-
imal region of pollex of propodus of major
cheliped of adult with large, conspicuously
flattened tooth. Exorbital angle tooth large,
triangular; intermediate tooth large, triangu-
lar blunt, as big as exorbital angle tooth. Epi-
branchial tooth small, about half size of in-
termediate tooth and exorbital angle tooth.
Anterolateral margin behind epibranchial
tooth smooth. Terminal article of gonopod 1
thin, needle-like, subterminal segment of
gonopod 1 slim. This is the largest species
of freshwater crab in Africa. Adult sizes
range from the size at the pubertal molt (cw
70—75 mm) to largest the known specimen
(cw 113 mm).
Description.—For a detailed description
see Cumberlidge (1995a, 1999). For a brief
description of the type see Capart (1954,
figs: 1546):
Remarks.—Sudanonautes africanus is a
common and well-known species that was
recently redescribed (Cumberlidge 1995a,
1999):
Ecology.—This species is restricted to the
more humid areas of the coastal rainforest
belt from south-east Nigeria to the mouth of
the Zaire river. Sudanonautes africanus oc-
curs in a range of permanent aquatic habitats
from large rivers and small streams (with
both fast and slow flowing water) to ponds.
In the Central African Republic S. africanus
is found in creeks up to 1.5 m deep with a
VOLUME 113, NUMBER 2
sand, gravel or mud bottom. Specimens
were also taken from burrows in overhang-
ing banks, among roots and under logs. Else-
where in its range, this species is also com-
mon in streams and rivers draining mature
forest, and has been reported to dig burrows
near waterways. This crab also occurs in
temporary water sources such as drainage
culverts and ditches. For more details see
Cumberlidge (1995a, 1999).
Distribution.—Sudanonautes africanus
occurs in the coastal rainforest regions of Ni-
geria and Central Africa. In Central Africa
S. africanus occurs in south Cameroon, the
Republic Populaire du Congo, and Gabon
(in the San Benito, Ogoué and Alima rivers),
and in the lower reaches of the Zaire River
basin. For more details see Cumberlidge
(1995a, 1999). The present record is the first
report of the presence of S. africanus in the
Central African Republic.
Sudanonautes sangha, new species
Fig. 4
Type material and type locality.—Central
African Republic. Holotype: 1 subadult fe-
male, cw 40.5, cl 29.7, ch 12.5, fw 10.7 mm
(AMNH 17825), a few km upstream from
the village of Bayanga (02°45'43’N,
16°14’12"E), Sangha river, depth 1—2 m, in
fish trap, eating worm, coll. J. Sullivan and
J. B. Kindimoungo, 18 Jun 1998.
Diagnosis.—Exorbital tooth large, point-
ed; intermediate tooth small, low; epibran-
chial tooth small, low, not directed outward,
set back behind mid-point of postfrontal
crest. Postfrontal crest spanning entire cara-
pace, crest curving backward before meeting
epibranchial tooth, anterolateral margin pos-
terior to epibranchial tooth raised, lined by
small granules. Semi-circular, urogastric,
transverse branchial grooves very deep. Ver-
tical suture on carapace sidewall meeting in-
termediate tooth. Carapace medium height
(ch/fw = 1.16). Mandibular palp two-seg-
mented; terminal segment single, undivided,
with hair at junction between segments. First
carpal tooth on carpus of cheliped large,
415
pointed; second carpal tooth reduced to
small granule.
Description.—Carapace (Figs. 4A, B).—
Ovoid, widest in anterior third (cw/fw 3.79),
medium height (ch/fw = 1.16), semi-circu-
lar, urogastric, transverse branchial grooves
very deep, regions smooth; cardiac region
weakly marked, cervical grooves present but
weak. Front slightly bilobed, anterior margin
indented, relatively narrow, about one-quar-
ter carapace width (fw/cw = 0.26) (Fig. 4B).
Postfrontal crest smooth, spanning entire
carapace, straight part consisting of fused
epigastric, postorbital crests, then curving
backward behind intermediate teeth to meet
anterolateral margins at epibranchial teeth.
Anterolateral margin smooth posterior to
epibranchial tooth. Exorbital tooth large,
sharp, pointed forward. Epibranchial tooth
low, small, set back behind mid-point of
postfrontal crest.
Carapace sidewalls mostly smooth, with
faint granules in suborbital regions. Each
sidewall with two sutures, one longitudinal,
one vertical, dividing sidewall into three
parts. Longitudinal (epimeral) suture divid-
ing suborbital, subhepatic regions from pter-
ygostomial region, beginning medially at
lower margin of orbit, curving backward
across flank. Short vertical suture dividing
suborbital region from subhepatic region;
vertical suture meeting intermediate tooth.
First transverse groove on sternum, between
sternal segments s2 and s3, complete; sec-
ond groove, between sternal segments s3
and s4, consisting of two small notches at
sides of sternum. Third maxillipeds filling
entire oral field, except for transversely oval
efferent respiratory openings at superior lat-
eral corners; long flagellum on exopod of
third maxilliped; ischium of third maxilliped
smooth, with clear vertical groove (Fig. 4C).
Mandibular palp two-segmented; terminal
segment single, undivided, with hair but no
hard flap at junction between segments (Fig.
4D). Segments 1—6 of female abdomen four-
sided, last segment a broad rounded triangle,
sides forming a smooth curved, rounded
margin; segments 5—6 broadest (Fig. 4E).
416 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Sudanonautes sangha, new species. Holotype subadult female, cw 40.5 mm, AMNH 17825. A,
carapace and eyes, dorsal view; B, cephalothorax, carapace and eyes, frontal view; C, left third maxilliped; D,
left mandible; E, abdomen; FE right cheliped, frontal view; G, left cheliped, frontal view; H, carpus and merus
of right cheliped, lateral view; I, carpus and merus of right cheliped, mesial view; J, left second pereiopod,
lateral view. Scale = 4.4 mm (D), 6.2 mm (I), 6.7 mm (C), 7.2 mm (H), 8.6 mm (EF G, J), and 12.7 mm (A,
B, E).
Dactylus of right cheliped long, slim, granules, cluster of granules surrounding
straight; palm of propodus swollen; fingers larger pointed distal meral tooth at distal end
of digits of chelipeds with small even teeth, (Fig. 41). Inner margin of carpus of cheliped
forming long slim interspace when closed, with large, slender, pointed tooth (first carpal
fingers almost meeting when shut (Fig. 4F). tooth), second carpal tooth reduced to a
Inferior margins of merus with rows of small granule (Fig. 41). Pereiopods P2—P5 slender
VOLUME 113, NUMBER 2
(Fig. 4J), P4 longest, P5 shortest. Propodus
of P2—P5 broad, posterior margin of propo-
dus of P2—P5 serrated, dactyli of P2—P5 ta-
pering to point, each bearing four rows of
downward-pointing sharp bristles; dactylus
of P5 shortest.
Etymology.—tThe species is named for the
Sangha river where it was collected. The
Sangha river drains a large region of the
Central African Republic, and is a dominant
natural feature of the area. The species name
sangha is a noun in apposition.
Remarks.—It is not normally good prac-
tice to describe a new species from a sub-
adult female. However, we have decided to
establish this taxon in light of the distinct
nature of the available morphological char-
acters, and because of the isolated nature of
the study area which may mean that further
specimens are unlikely to become available
for some time. Sudanonautes sangha is the
eleventh species of this West and Central Af-
rican genus. Characters of the gonopods,
adult male chelipeds, abdomen and sternum
are not available because the only specimen
of S. sangha is a subadult female. Neverthe-
less, there are a number of unique characters
that distinguish S. sangha from other species
in the genus.
Sudanonautes sangha is most likely to be
confused with other large species of fresh-
water crabs occurring in the rain forests of
Central Africa such as S$. chavanesii, S. far-
adjensis, S. africanus, S. aubryi and S. flow-
Crt:
Sudanonautes sangha and S. africanus are
similar in that both species have a small epi-
branchial tooth, and both lack large teeth on
the anterolateral margins of the carapace.
However, there are a number of characters
which distinguish S. sangha from S. african-
us. The carapace of S. sangha is not as flat-
tened as that of S. africanus (ch/fw S. san-
gha = 1.16, S. africanus = 1.06), and the
posterior region of the carapace of S. sangha
is smooth, whereas that of S. africanus is
rough with warty patches and ridges in the
posterior region. The cardiac regions of S.
sangha are flattened and are neither rounded
417
nor well marked, whereas these regions in
S. africanus appear as a pair of distinct
raised rounded structures that are clearly
outlined by deep cardiac grooves. The inter-
mediate tooth on the anterolateral margin be-
tween the exorbital tooth and the epibran-
chial tooth of S. sangha is small and low,
whereas that of S. africanus is large and tri-
angular. The second carpal tooth on the che-
liped of S. sangha is very small, only the
size of a granule, whereas the second carpal
tooth of S. africanus, while smaller than the
first carpal tooth, is distinct, pointed, and is
a tooth rather than a granule. The vertical
groove on the carapace sidewall of S. san-
gha meets the anterolateral margin at the
base of the intermediate tooth, whereas the
vertical groove of S. africanus meets the an-
terolateral margin at the base of the epibran-
chial tooth.
Sudanonautes sangha can be distin-
guished from S. chavanesii by the position
of the postfrontal crest and by the shape of
the epibranchial teeth. In S. sangha the lat-
eral ends of the postfrontal crest curve for-
ward to meet the epibranchial teeth in line
with the mid groove of the crest, whereas in
S. chavanesii the lateral ends of the post-
frontal crest curve sharply backward before
meeting the epibranchial teeth which are set
back posterior to the mid groove of the crest.
In S$. sangha, the epibranchial tooth is small,
blunt, pointed forward, and positioned in
line with the mid groove of the crest, where-
as in S. chavanesii, the epibranchial tooth is
large, sharp, pointed outward, and positioned
well behind the mid-groove of the postfron-
tal crest. Further, in S. sangha the vertical
suture on the carapace sidewall is simple,
whereas in S. chavanesii the vertical suture
on the carapace sidewall forms a Y-shaped
depression beneath the intermediate tooth.
Finally, in S. sangha the junction between
the two segments of the mandibular palp is
simple, whereas in S. chavanesii there is a
small hard flap on the mandibular palp at the
junction between the two segments.
Sudanonautes sangha can be distin-
guished from S. faradjensis by the form of
418
the anterolateral margin behind the epibran-
chial tooth: that of S. sangha is raised and
lined by small granules, whereas in S. far-
adjensis, there 1s a row of sharp teeth. Su-
danonautes sangha can be distinguished
from S. granulatus as follows: the exorbital
tooth of S. sangha is wide and triangular,
whereas that of S. granulatus is narrow and
low, and the anterolateral margin of S. san-
gha is raised and lined by granules, whereas
that of S. granulatus is completely smooth.
Sudanonautes sangha can be distin-
guished from S. aubryi by differences in the
postfrontal crest: that of S. sangha meets the
anterolateral margin at the epibranchial
tooth, whereas that of S. aubryi meets the
anterolateral margin behind the epibranchial
tooth. In addition, the carapace of S. sangha
is distinctly flatter than that of S. aubryi (ch/
fw S. sangha = 1.16, S. aubryi = 1.29).
Finally, S. sangha can be distinguished
from S. floweri by differences in the man-
dibular palp: in S. sangha the junction be-
tween the two segments of the mandibular
palp is simple, whereas in S. floweri there is
a small hard flap on the mandibular palp at
the junction between the two segments. In
addition, the carapace of S. sangha is dis-
tinctly flatter than that of S. floweri (ch/fw
S. sangha = 1.16, S. floweri = 1.68).
Ecology.—Collected in a large river (as
opposed to a small stream), and caught in a
baited trap.
Distribution.—Central African Republic,
a few kilometers upstream from the village
of Bayanga (02°45'43"N, 16°14'12”E), San-
gha river.
Acknowledgments
Marie Lawrence and James Cordeiro
(AMNH) are thanked for making this ma-
terial available to the authors for identifica-
tion. Alan Harvey (Georgia Southern Uni-
versity) provided training in computer aided
biological illustration to CBB. Collection of
this material was made possible through the
efforts of the AMNH Center for Biodiversity
Conservation and the World Wildlife Fund.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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Man, J. G., de. 1901. Description of a new fresh-water
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. 1903. On Potamon (Potamonautes) latidactyl-
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um, a new fresh-water crab from Upper Guin-
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Milne-Edwards, A. 1869. Révision du genre Thelphusa
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. 1886. La description de quelques Crustacés du
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les régions du Congo.—Bulletin de la Société
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. 1887. Observations sur les crabes des eaux
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Milne Edwards, H. 1853. Mémoire sur la famille des
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Monod, T. 1977. Sur quelques crustacés Décapodes af-
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. 1980. Décapodes. Pp. 369-389. Ed. J. R. Du-
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1905. Les crabes d’eau douce (Potamoni-
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. 1921. The brachyuran crabs collected by the
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Roux, J. 1927. Note sur une collection de Crustacés
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Stewart, B. A. 1997a. Morphological and genetic dif-
ferentiation between populations of river crabs
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. 1997b. Biochemical and morphological evi-
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Stewart, B. A., M. Coke, & P A. Cook, 1995. Pota-
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):420—425. 2000.
The freshwater crabs of the Barbilla National Park, Costa Rica
(Crustacea: Brachyura: Pseudothelphusidae), with notes on the
evolution of structures for spermatophore retention
Gilberto Rodriguez and Ingemar Hedstr6m
(GR) Centro de Ecologia, Instituto Venezolano de Investigaciones Cientificas, Apartado 21827,
Caracas 1020 A, Venezuela, email:grodrigu @ oikos.ivic.ve; (IH) Mid Sweden University,
Department of Applied Science, S-871 88 Harnésand, Sweden, email:
ingemar.hedstrom @tnv.mh.se.
Abstract.—The Barbilla National Park, a natural area of high biodiversity
on the Caribbean slope of Costa Rica, possesses two species of freshwater
crabs that share the same biotope in different localities of the Park. The first
species, Potamocarcinus magnus (Rathbun, 1896), one of the largest species
of the family Pseudothelphusidae, is widely distributed in Middle America,
from Costa Rica to Southern Mexico. The other species is a new species,
Ptychophallus barbillaensis. This is a species of small crabs, possibly restricted
to the National Park and neighboring areas. It can be distinguished from all
other species in the genus by the form of the receptacle formed in the apex of
the male gonopod, possibly for the retention of spermatophora during copu-
lation. The species of Ptychophallus Smalley, can be arranged in a morphocline
according to the relative development of this receptacle, with the present new
species midway between the ancestral condition and the closed channel found
in P. goldmanni Pretzmann, 1965.
The Barbilla National Park covers 12,830
hectares of humid tropical forest on the Ca-
ribbean watershed of the Sierra de Tala-
manca, Costa Rica. This stretch of mostly
primary vegetation is located between the
valleys of the Pacuare and Chirrip6o rivers,
between 200 and 1600 m above sea level.
The area is noted for its high biodiversity.
During recent surveys carried out by the
Mid Sweden University several samples of
freshwater organisms have been collected
in the ravines that surround the Nairi Field
Station, close to the northwestern border of
the Park. Two species of freshwater crabs
were discovered among the materials in
these collections. One of them, a new spe-
cies belonging to the genus Ptychophallus
Smalley, 1964a, is described in the present
contribution. The materials recorded are de-
posited in the Reference Collection of the
Instituto Venezolano de Investigaciones
Cientificas, Caracas (IVIC); the National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. (USNM), and
the Museum of Natural History of Tulane
University, New Orleans (TU). Other ab-
breviations used are cl, carapace length, and
cb, carapace breadth.
Systematics
Family Pseudothelphusidae Rathbun, 1893
Genus Potamocarcinus H. Milne Edwards,
1555
Potamocarcinus magnus (Rathbun, 1896)
Material.—Costa Rica: Rio Cano Seco
(10°00’N, 83°26'W), affluent of Rio Dantas,
Barbilla National Park, 7 Jan 1999, leg. I.
Hedstrom, 300 m above sea level, 1 im-
mature female, cl 17.3 mm, cb 25.0 mm
VOLUME 113, NUMBER 2
(IVIC 1070); Casas Negras, Rio Dantas,
Barbilla National Park, 8 Jan 1999, leg. I.
Hedstrém, 200 m above sea level, 1 im-
mature male, cl 19.8 mm, cb 28.3 mm
(IVIC 1071); Rio Barbilla, Barbilla Nation-
al Park, 28 Jan 1999, leg. I. Hedstr6m, 100
m above sea level, 2 immature males, cl
17.0 and 12.4 mm, cb 23.7 and 18.5 mm,
1 immature female, cl 10.0 mm, cb 15.7
mm (IVIC 1072).
Remarks.—Although our specimens are
immature, they can be referred with cer-
tainty to this species. Potamocarcinus mag-
nus 1s widely distributed in Central Amer-
ica, but Rodriguez (1982) distinguish three
different morphological groups from (a)
Costa Rica, (b) El Salvador, Guatemala and
southern Mexico, and (c) Guatemala. The
characters of the carapace in our specimens
agree with those given for the Costarican
form. However, the typical flat tubercles
present over the surface of the carapace
cover a greater area, a characteristic possi-
bly due to the juvenile condition of the
specimens. The right cheliped has the typ-
ical long and narrow tubercle over the
palm, at the base of the fingers, although it
is relatively litthe developed. In the first
male gonopods there is not an abnormal in-
trusion of the marginal process into the dis-
tal lobe, as was the case in the specimen
from Costa Rica illustrated by Rodriguez
(1982, fig. 72b). Potamocarcinus magnus is
the largest of all species of Pseudothelphu-
sidae, with a cb of 135 mm reported by
Rathbun (1896).
Genus Ptychophallus Smalley, 1964a
Ptychophallus barbillaensis, new species
Pig. 1 2C)
Material.—Costa Rica: Rio Cano Seco
(10°00’N; 83°26’W), affluent of Rio Dantas,
Barbilla National Park, 7 Jan 1999, leg. I.
Hedstr6m, 300 m above sea level, 1 male
holotype, cl 16.9 mm, cb 27.7 mm (IVIC
1073), 1 male paratype, cl 13.4 mm, cb 20.7
mm (IVIC 1074); Nairi Field Station
(09°59'N; 83°27'W), 3 km north of the bor-
421
der of Barbilla National Park, 7 Jan 1999,
leg. I. Hedstrém, 2 ovigerous females, cl
15.6 and 13.9 mm, cb 25.0 and 21.1 mm
(IVIC 1075); Casas Negras, Rio Dantas,
Barbilla National Park, 8 Jan 1999, leg. I.
Hedstr6m, 200 m above sea level, 2 males,
cl 12.2 and 11.9 mm, cb 18.9 and 18.7 mm
(IVIC 1076); Las Cuevas, Rio Dantas, Bar-
billa National Park, 11 Jan 1999, leg. I.
Hedstrom, 150 m above sea level, 2 males,
cl 14.2 and 11.7 mm, cb 22.5 and 18.6 mm,
1 mature female, cl 14.1 mm, cb 21.5 mm
(IVIC 1077); El Recodo, Rio Dantas, Bar-
billa National Park, 6 Jan 1999, leg. I. Hed-
strom, 300 m above sea level, 1 male, cl
15.2 mm, cb 23.3 mm, 1 immature male, cl
9.9 mm, cb 15.0 mm, 1 male juvenile, cl
5.8 mm, cb 13.5 mm (IVIC 1078).
Diagnosis.—First gonopods with large
lateral lobe divided in 2 subequal rounded
segments by median notch; proximal seg-
ment smaller, subcircular; distal segment
projected anteriorly and bent caudally to
form cup-shaped receptacle; apex strongly
bent laterally forming, with distal segment
of lateral lobe, characteristic sinus; field of
spines directed toward latero-cephalic side,
oblong, with deep notch on mesial side; ce-
phalic end of apex with 2 expansions, distal
subtriangular in lateral and caudal views,
with notch on lateral side, proximal round-
ed in cephalic view, beak-like in mesial
view.
Description of holotype.—Carapace 1.6
times as wide as long, surface smooth and
polished, except for few granules on pos-
terior branchial regions, near margins; cer-
vical grooves recurved backwards, narrow
and deep, not reaching margins of carapace;
anterolateral margins with shallow and
wide postorbital notch bordered by 5 papil-
lae, rest of borders with small papillae that
becomes dentiform behind level of cervical
grooves. Postfrontal lobes low, delimited
anteriorly by transverse depressions; medi-
an groove narrow, deep, making incision on
upper margin of front. Surface of carapace
between postfrontal lobes and front flat,
slightly inclined forward and towards mid-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figw le
Ptychophallus barbillaensis, new species, holotype from Rio Cafio Seco, Costa Rica (IVIC 1073):
A, dorsal view of right side of carapace; B, third maxilliped, left; C, chela of largest cheliped (left), external
view; D, first left gonopod, caudal view; E, same, cephalic view; EK same, apex, mesial view; G, same, apex,
cephalic view.
dle line. Upper margin of front in dorsal
view slightly convex and divided into two
halves by median notch, thin, well marked,
with small tubercles; lower margin thin,
moderately sinuous, advanced in front of
upper margin; both margins subparallel.
Exognath of third maxilliped 0.65 length
of ischium of endognath. Palm of largest
cheliped moderately swollen, with lower
and upper margins convex; fingers gaping,
with rows of small black-brown points on
external surface.
First gonopods wide in latero-mesial di-
rection, narrow in caudo-cephalic direction;
large lateral lobe divided in 2 subequal
rounded segments by median notch; proxi-
mal segment smaller, subcircular; distal
segment projected anteriorly and bent cau-
dally to form cup-shaped receptacle semi-
enclosed by apex of gonopod; apex strongly
VOLUME 113, NUMBER 2
bent laterally, forming with distal segment
of lateral lobe characteristic sinus; field of
spines directed toward latero-cephalic side,
oblong, with deep notch on mesial side; ce-
phalic end of apex forming two expansions;
distal expansion continuous with margin of
gonopod, subtriangular, in lateral and cau-
dal views, with notch on lateral side; prox-
imal expansion rounded in cephalic view,
beak-like in mesial view.
Remarks.—This species can be distin-
guished from all others in the genus by the
cupshaped expansion of the distal segment
of the lateral lobe of the first gonopods and
by the strong recurvation of the apical pe-
duncle of gonopod over this expansion.
Etymology.—The species is named after
the Barbilla National Park, Costa Rica,
where the species was collected.
Evolution of Structures for Spermatophore
Retention in Ptychophallus
Although the morphology of the first
male gonopods are essential for the dis-
crimination of the species of Pseudothel-
phusidae, very little is known of the func-
tion of the different processes of these ap-
pendages during copulation. In all species,
the apex is provided with a field of minute
spines (Smalley 1964b) which surround the
opening of the spermatic channel and serve
to hold the spermatophore once it is extrud-
ed by the piston-like pumping action of the
second gonopod.
In the species of Ptychophallus a trend
can be observed towards a reaccomodation
of the lateral lobe and the various compo-
nents of the apex of the gonopod, to form
a structure that possibly serves to receive
and keep the spermatophore once it is ex-
truded. This is fully developed in Ptycho-
Phallus goldmanni Pretzmann, 1965, but
the other species can be arranged in a mor-
phocline comprising several stages of de-
velopment of this structure, as follows.
Stage I (Fig. 2A, F).—In the primitive
condition there is a wide lateral lobe, which
is foliose, with a shallow depression or
423
deeply notched, with the distal portion ad-
vanced and transverse. The field of spines
is located distant from the lateral lobe, fac-
ing caudo-laterally, its long axis forming an
angle of approximately 60° with the longi-
tudinal axis of the appendage. Species at
this stage are P. colombianus (Rathbun,
1893), P. exillipes (Rathbun, 1898), P. kuna
Campos & Lemaitre, 1999, and P. tristani
(Rathbun, 1896).
Stage II (Fig. 2B, G).—Lateral lobe
deeply notched as in some species of stage
I, but he distal portion is more advanced
and joins the caudal side of the apical pe-
duncle. The field of spines approaches the
lateral lobe and is directed laterally, with its
long axis forming an angle of approximate-
ly 180° with the longitudinal axis of the ap-
pendage. Species at this stage are P. micra-
canthus Rodriguez, 1994, P. montanus
(Rathbun, 1898), and P. tumimanus (Rath-
bun, 1898).
Stage IIIT (Fig. 2C, H).—Lateral lobe as
in stage II, but the distal portion is cupped,
and its border is continuous with the caudal
margin of the apical peduncle. The field of
spines faces the lateral side, and its long
axis forms an angle of approximately 230°
with the longitudinal axis of the appendage.
The only species at this stage is P. barbil-
laensis.
Stage IV (Fig. 2D, I).—The lateral lobe
has the proximal segment strongly reduced,
the distal one very advanced and forms, to-
gether with the border of the field of spines,
a channel-like receptacle which is wide
open. The field of spines is directed later-
ally, its long axis forming an angle of ap-
proximately 180° with the longitudinal axis
of the appendage. The only species at this
stage is P. cocleensis Pretzmann, 1965.
Stage V (Fig. 2E, J).—The lateral lobe is
reduced to a distal segment, very advanced
and forms, together with the border of the
field of spines, an almost closed channel-
like receptacle. The field of spines is di-
rected laterally, its long axis forming an an-
gle of approximately 180° with the longi-
tudinal axis of appendage. The only species
424
Pier 2.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
a-e 2mm
iL
First left gonopod of Ptychophallus species: A, EF P. tristani (Rathbun, 1896), male from ~1.8 km
E Atenas, San José Province, Costa Rica (TU 4436); B, G, P. montanus (Rathbun, 1898), male from ~3 km N
San Jer6nimo de Moravia, San José Province, Costa Rica (TU 4442); C, H, P. barbillaensis, new species,
holotype from Rio Cafio Seco, Costa Rica (IVIC 1073); D, I, P. cocleensis Pretzmann, 1965, holotype from Rio
Coclé del Norte, Panama (USNM 119869); E, J, P. goldmanni Pretzmann, 1965, holotype from Cana, Panama
(USNM 54044); A-E, detail of apex, lateral view: F—J, total view, caudal.
at this stage is P. goldmanni Pretzmann,
1965.
A similar morphocline, although not
leading to the formation of a structure for
the retention of the spermatophore, has
been observed in the lateral lobe of species
of the genera Pseudothelphusa Saussure,
1857, and Tehuana Rodriguez & Smalley,
1969, in southern Mexico (Rodriguez
1986). In this case the morphocline follows
a general westward direction, suggesting
an allopatric speciation of primitive demes,
encompassing a migration along the same
geographical axis. In the case of Ptycho-
phallus the pattern is more complex. Spe-
cies of stages I and II overlap their areas
of distribution in Costa Rica, and the areas
of P. kuna (stage I) and P. micracanthus
(stage II) are relatively close in Central
Panama (see map in Campos & Lemaitre
1999). The area of P. barbillaensis (stage
III) is nested among the species of stages
I and II. P. cocleensis (Stage IV) is found
in Central Panama, and P. goldmanni
(Stage IV) further East, near the Colom-
bian border. Notwithstanding this complex
pattern, the general trend of the morphoc-
line is eastward. This direction contrasts
with the supposed radiation of the family,
that proceeded westward from an ancestral
area in northern Colombia (Rodriguez
1986).
Two species of Ptychophallus cannot be
placed in this morphocline. P. lavallensis
Pretzmann, 1978, has a field of spines
strongly upturned cephalically and a very
wide undivided lateral lobe that does not
approach the field of spines. P. paraxan-
thusi Bott, 1968, has a field of spines bent
mesially and a wide, shallow-notched, lat-
eral lobe that displays distally several ridg-
es but does not form a receptacle. These
species possibly represent phyletic lines
that differ both among themselves, as with
those species in the morphocline described
above.
VOLUME 113, NUMBER 2
Literature Cited
Bott, R. 1968. Fluss-Krabben aus dem 6stlichen Mittel-
Amerika und von den Grossen Antillen (Crus-
tacea, Decapoda).—Senckenbergiana Biologica
49:39—49.
Campos, M. R., & R. Lemaitre. 1999. Two new fresh-
water crabs of the genus Ptychophallus Smal-
ley, 1964 (Crustacea: Decapoda: Brachyura:
Pseudothelphusidae) from Panama, with notes
on the distribution of the genus.—Proceedings
of the Biological Society of Washington 112:
553-561.
Milne Edwards, H. 1853. Memoire sur la famille des
Ocypodien.—Annales des Sciences Naturelles,
Zoologie, 3e série 20:163—228.
Pretzmann, G. 1965. Vorlaufiger Bericht tiber die Fam-
ilie Pseudothelphusidae.—Anzeiger der Mathe-
matisch Naturwissenschaftliche Klasse der Os-
terreichischen Akademie der Wissenschaften (1)
1:1-10.
. 1978. Neue Potamocarcinini, Poglayen-Neu-
wall leg 1975 (Vorlaufige Mitteilung).—Sit-
zungsberichten der Osterreichischen Akademie
der Wissenschaften, Mathematisch-Naturwis-
senschaftliche Klasse (1) 1978 (2):51—54.
Rathbun, M. J. 1893. Descriptions of new species of
American freshwater crabs.—Proceedings of
the United States National Museum 16(959):
649-661.
. 1896. Descriptions of two species of fresh-
water crabs from Costa Rica.—Proceedings of
425
the United States National Museum 18(1071):
377-379, fig. 1-3, pl. 29-30.
. 1898. A contribution to a knowledge of the
freshwater crabs of America. The Pseudothel-
phusinae.—Proceedings of the United States
National Museum 21(1158):507—537.
Rodriguez, G. 1982. Les crabes d’eau douce
d’Amérique. Famille des Pseudothelphusi-
dae.—Faune Tropicale 22:1—223.
. Centers of distribution of Neotropical fresh-
water crabs. Jn R. H. Gore & K. L. Heck, eds.,
Biogeography of the Crustacea.—Crustacean
Issues 3:51—67.
. 1994. A revision of the type material of some
species of Hypolobocera and Ptychophallus
(Crustacea: Decapoda: Pseudothelphusidae) in
the National Museum of Natural History, Wash-
ington, D.C. with descriptions of a new species
and a new subspecies.—Proceedings of the Bi-
ological Society of Washington 107:296—307.
,& A. E. Smalley. 1969. Los cangrejos de agua
de Mexico.—Anales del Instituto de Biologia
(Mexico) 41:69—112.
Saussure, H. de. 1857. Diagnoses de quelques Crus-
tacés nouveaux des Antilles et du Mexique.—
Revue et Magazine de Zoologie Pure et Appli-
qué (2)9:304—306.
Smalley, A. E. 1964a. The river crabs of Costa Rica,
and the subfamilies of the Pseudothelphusi-
dae.—Tulane Studies in Zoology 12:5-—13.
. 1964b. A terminology for the gonopods of the
American river crabs.—Systematic Zoology 13:
28-31.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):426—430. 2000.
Acanthilia, a new genus of leucosioid crabs (Crustacea: Brachyura)
from the Atlantic coast of the Americas
Bella S. Galil
National Institute of Oceanography, Israel Oceanographic & Limnological Research, P.O.B. 8030,
Haifa 31080, Israel
Abstract.—A new monotypic leucosioid genus, Acanthilia, is established
from the Atlantic coast of the Americas, for one species of the genus /liacantha
Stimpson, J. intermedia Miers. The new genus differs from /liacantha in its
tridentate anterior margin of the efferent branchial channel, the shorter cheliped
fingers, the glabrous pereiopodal dactyls, the convex margins of the sixth ab-
dominal segment in the male, and the claw-tipped first male pleopod.
As part of an on-going study of leuco-
sioidean crabs, the leucosioids of the Atlan-
tic coast of the Americas at the National
Museum of Natural History, Smithsonian
Institution (USNM), were examined. An in-
vestigation of /liacantha Stimpson, 1871,
showed that J. intermedia Miers, 1886, dif-
fers from the five other members of the ge-
nus in its tridentate anterior margin of the
efferent channel, the shorter cheliped fin-
gers, the glabrous pereiopodal dactyls, the
convex margins of the sixth abdominal seg-
ment in the male, and the claw-tipped male
pleopod. J. intermedia was removed from
the genus /liacantha and placed in a new
genus herein established.
Acanthilia, new genus
Diagnosis.—Carapace subovate, glo-
bose; regions indistinct. Front narrow,
prominent, bilobed. Basal antennular seg-
ment forming an operculum, partially seal-
ing antennular aperture. Antennular fossa
not separated from orbit. Orbital margin
with 3 sutures, basal antennal segment in-
serted in orbital hiatus proximally on ven-
tral margin. Postorbital region concave.
Third maxillipeds fitting closely, sealing
buccal cavity, efferent branchial channel.
Inner surface of 3 maxilliped exognath gla-
brous. Anterior margin of efferent branchial
channel produced, tridentate, contiguous
with lower orbital margin.
Anterolateral margin of carapace medi-
ally concave, posterolateral margin uni-
formly curved. Posterior margin of cara-
pace bidentate. Spine on posterior intestinal
region upcurved, projecting beyond poste-
rior denticles. Abdominal sulcus in male
deep, nearly reaching buccal cavity.
Chelipeds long; merus_ subcylindrical;
propodus swollen basally; fingers half as
long as propodus, opening vertically. Pe-
reiopods short, dactyls triqueterous, longer
than propodi, glabrous.
Male abdomen with segments 3—5 fused,
tapering distally, basio-lateral regions of
fused segments somewhat inflated; lateral
margins of segment 6 convex. Female ab-
domen with segments 4—6 fused, greatly
swollen.
Male first pleopod stout, tip claw-shaped;
second pleopod short, distally scoop-like.
Type species.—lIliacantha intermedia
Miers, 1886.
Etymology.—Acanthilia is an anagram of
Iliacantha Stimpson, 1871.
Remarks.—Stimpson (1871) erected the
genus Jliacantha for two species, I. subgl-
obosa and I. sparsa, collected off Florida,
and characterized by three posterior spines,
VOLUME 113, NUMBER 2 427
QoL28
Fig. 1. Acanthilia intermedia (Miers, 1886).—dé (USNM 274840). A, carapace and right cheliped, dorsal
view; B, anterior margin of efferent branchial channel; C, sixth abdominal segment and telson, ventral view; D,
tip of first pleopod, ventral view. A, 5 mm; B,C,D, 0.1 mm.
428 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Iliacantha subglobosa.—é (USNM 274724). A, carapace and right cheliped, dorsal view; B, anterior
margin of efferent branchial channel, ventral view; C, sixth abdominal segment and telson, ventral view; D, tip
of first pleopod, ventral view. A, 5 mm; B,C,D, 0.1 mm.
VOLUME 113, NUMBER 2
cheliped fingers opening vertically, and pro-
jecting pterygostomian channels. When de-
scribing J. intermedia, Miers (1886:302)
noted it differed from Stimpson’s species in
the “‘length of the dactyli of the chelipeds’’.
Acanthilia, new genus, is easily distin-
guished from J/liacantha by the tridentate
margin of the efferent branchial channel
(Fig. 1B), the cheliped fingers half as long
as the propodus (Fig. 1A), the glabrous per-
eiopodal dactyls, the convex margins of the
male abdominal segment 6 (Fig. 1C), and
the claw-tipped first male pleopod (Fig.
1D); whereas in /liacantha the margin of
efferent channel is bilobate (Fig. 2B), the
cheliped fingers are longer than the propo-
dus (Fig. 2A), pereiopodal dactyls are se-
tose, margins of the male abdominal seg-
ment 6 are straight (Fig. 2C), and the first
male pleopod is distally tapering, simple
(Fig. 2D).
Acanthilia intermedia (Miers, 1886)
Fig. 1
Iliacantha intermedia Miers, 1886:302, pl.
26, fig. 3—Hay & Shore, 1918:424, pl.
32, figs. 3, 3a.—Rathbun, 1937:186, pl.
54, figs. 1, 2.—Williams, 1965:151, fig.
129; 1984:290, fig. 225.—Coelho, 1970:
234.—Coelho & Ramos, 1972:184.—
Fausto-Filho, 1975:81; 1979:51.—G6-
mez & Ortiz, 1976:8.—Powers, 1977:
37.—Coelho & Torres, 1980:72.—Ro-
drigues, 1980:259.—Takeda, 1983:
118.—Keith, 1985:254, fig. 2f—Abele
& Kim, 1986:42, fig. 485 b.—Taissoun,
1988:126.—Werding & Miiller, 1990:
412, figs. 7a—c.—de Melo, 1996:159,
fig.—de Melo et al., 1998:450.
Iliacantha liodactylus Takeda,
fig.
1983:159;
Material examined.—United States.
North Carolina, 33°20’N, 77°46’W, 25 m,
16 Aug 1977, 2 66 15.8, 20:0mm, ¢ juv.
12.2 mm (USNM_ 174250).—Georgia,
31°41.12’N, 80°20.48’W, 28 m, 30 Jan
1980, ¢ 21.6 mm (USNM 214909).—Sa-
pelo Is., 22 m, 5 Mar 1963, coll. M. Gray,
429
6 22.5 mm (USNM_ 155484).—Florida,
25°45.56'N, 82°09.21'W, 19.6 m, 28 Apr
1981, coll. R. Lemaitre, ovigerous @ 25.1
mm (USNM 242750).
Redescription.—Dorsal surface of cara-
pace granulate, front minutely shagreened.
Fronto-orbital margin half as wide as pos-
terior margin, slightly concave. Anterolat-
eral margin set with prominent pearliform
granules, subhepatic margin somewhat in-
flated. Posterior margin bearing pair of dor-
so-ventrally flattened triangular denticles
laterally, pearliform granules medially. In-
testinal spine prominent, granulate, upcur-
ved.
Anterior margin of efferent branchial
channel prominently tridentate, advanced
beyond orbital margin. Third maxilliped
bearing fungiform granules, crowded ante-
riorly, female ischium longitudinally tra-
versed by row of setae.
Cheliped merus granulose, granules larg-
er proximally; carpus and propodus smooth,
fingers costate. Pereiopodal meri minutely
granulate, upper margin of propodi promi-
nently costate, dactyli styliform.
Thoracic sternites closely granulate.
Male abdomen minutely granulate, fused
segments of female abdomen smooth ex-
cept for granulate basal margin. Male first
pleopod slightly sinuous, dorso-ventrally
flattened, subapically setose; cornute tip
bearing subquadrate plate proximally, claw-
like appendix distally.
Color.—“‘gray, without markings of any
kind”’ (Hay & Shore 1918:424), ‘“‘Amarillo
crema O gris con puntos pequefnios dispersos
de color marr6n claro”? (Taissoun 1988:
126).
Remarks.—The color plates of J. inter-
media and I. liodactylus (Takeda, 1983:118,
119) were erroneously exchanged.
Distribution.—North Carolina to Brazil;
10—329 m.
Acknowledgements
I am deeply grateful to R. B. Manning
for the invitation to study the leucosioids in
430
the National Museum of Natural History,
Smithsonian Institution, Washington D.C.
The visit was supported by the Museum’s
Collection Improvement funds.
Literature Cited
Abele, L. G., & W. Kim. 1986. An illustrated guide to
the marine decapod crustaceans of Florida.
State of Florida Department of Environmental
Regulation, Tallahassee, 760 pp.
Coelho, P. A. 1970. A distribuigao dos crustaceos de-
capodos reptantes do norte do Brasil.—Trabal-
hos do Instituto Oceanograficos, Universidade
Federal de Pernambuco, Recife, 9/1 1:223—238.
, & M. A. Ramos. 1972. A constituigao e a
distribuigao da fauna de decapodos do litoral
leste da América do sul entre as latitudes de 5°
N e 39° S.—Trabalhos do Instituto Oceanograf-
icos, Universidade Federal de Pernambuco, Re-
cies 132135—2 36:
, & M. FE A. Torres. 1980. Zoogeografia mar-
inha do Brazil II. Consideragdes ecolégicas e
biogeograficas sobre a familia Leucosiidae (De-
capoda: Brachyura).—Revista nordestina de
biologia 3(especial):67—77.
Fausto-Filho, J. 1975. Quinta contribui¢gao ao inven-
tario dos crustaceos decapodos dos substratos
de cascalho do Nordeste brasileiro.—Ciencia
Agronomica 10:109—124.
. 1979. Crustaceos estomat6podos e decapodos
dos substratos de areia do nordeste brasileiro.—
Arquivos de Ciencias do Mar 19:45—56.
Gomez, O., & M. Ortiz. 1976. Lista de braquiuros cu-
banos.—Ciencias, Investigaciones Marinas,
Universidad de la Habana, 25:3—20.
Hay, W. P, & C. A. Shore. 1918. The decapod crus-
taceans of Beaufort, N.C. and surrounding re-
gion.—Bulletin of the United States Bureau of
Fisheries 35(1915/1916):369—475.
Keith, D. E. 1985. Shallow-water and terrestrial Brach-
yuran crabs of Roatan and the Swan Islands,
Honduras.—Sarsia 70:25 1-278.
Melo, G. A. S., de. 1996. Manual de identificacgao dos
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Brachyura (caranguejos e siris) do litoral bras-
ileiro. Pléiade/FAPESP, 604 pp.
, M. EF A. Torres, & O. Campos. 1998. Mala-
costraca-Eucarida. Brachyura. Dromiacea and
Oxystomata. Pp. 439—454 in PS. Young, Cata-
logue of Crustacea of Brazil. Rio de Janeiro
Museu Nacional.
Miers, E. J. 1886. Report on the Brachyura collected
by H.M.S. Challenger during the years 1873—
1876.—Report on the scientific results of the
voyage of H.M.S. “Challenger”, Zoology
17(49): 1-362.
Powers, L. W. 1977. A catalogue and bibliography to
the crabs (Brachyura) of the Gulf of Mexico.—
Contributions in Marine Science 20(supple-
ment):1—190.
Rathbun, M. J. 1937. The oxystomatous and allied
crabs of America.—United States National Mu-
seum Bulletin 166:1—278, 86 pls.
Rodriguez, G. 1980. Los Crustaceos decadpodos de
Venezuela. Instituto Venezolano de Investiga-
ciones Cientificas, Caracas, 444 pp., 70 figs.
Stimpson, W. 1871. Preliminary report on the Crusta-
cea dredged in the Gulf Stream in the Straits of
Florida, by L.E de Pourtales, Assist. U. S. Coast
Survey. Part I. Brachyura.—Bulletin of the Mu-
seum of Comparative Zoology 2:109—160.
Taissoun, N. E. 1986-88. Los cangrejos decapodos
Brachyura de las costas da Venezuela. III. Fam-
ilias Leucosiidae Dana, 1852 y Geryonidae
(Beurlen, 1930).—Boletin del Centro de Inves-
tigaciones Bioldgicas, Universidad del Zulia,
Maracaibo, 17:121—140.
Takeda, M. 1983. Crustaceans. In M. Takeda, & T.
Okutani, Crustaceans and mollusks trawled off
Suriname and French Guiana. Japan Marine
Fishery Resource Research Center, Tokyo, 185
PP.
Werding, B., & H. G. Miiller. 1990. Leucosiidae von
der nordkuste Kolumbiens (Crustacea: Deca-
poda: Brachyura).—Senckenbergiana Biologica
70:405—417.
Williams, A. B. 1965. Marine decapod crustaceans of
the Carolinas.—Fishery Bulletin 65:298 pp.
. 1984. Shrimps, lobsters, and crabs of the At-
lantic coast of the eastern United States, Maine
to Florida. Smithsonian Institution Press, Wash-
ington, D.C., 550 pp.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):431—442. 2000.
Cambarus (Cambarus) davidi, a new species of crayfish
(Decapoda: Cambaridae) from North Carolina
John E. Cooper
North Carolina State Museum of Natural Sciences, Research Lab, 4301 Reedy Creek Road,
Raleigh, North Carolina 27607, U.S.A.
Abstract.—Cambarus (Cambarus) davidi is a new species of crayfish from
the eastern Piedmont Plateau of North Carolina, where it is restricted to inter-
mittent streams, seepage areas, springs, and burrows. Although the ranges of
the two species appear to be broadly disjunct, C. (C.) davidi is most closely
related to the ecologically more tolerant Cambarus (C.) bartonii (s.1.), and
especially resembles some members of the controversial subspecies C. (C.) b.
cavatus. Cambarus (C.) davidi is distinguished by a suite of characters that
includes a vaulted carapace; a deeply excavate, ladlelike rostrum; a very nar-
row, sparsely punctate areola; and an obtuse to nearly obsolete suborbital angle
that almost always bears a small tubercle.
On 18 August 1993, David G. Cooper
collected several specimens of a Cambarus
from under large rocks in a shallow, inter-
mittent tributary of the Neuse River, Wake
County, North Carolina. In the field they
appeared to be aberrant individuals of the
burrowing species, Cambarus (Depressi-
cambarus) reduncus Hobbs, 1956, which is
not uncommon in such habitats in the upper
Neuse River watershed. In the laboratory,
however, I was surprised to discover that
the specimens belonged to the subgenus
Cambarus, and to some species previously
unknown from the Neuse River basin,
whose crayfish fauna is well documented
(Cooper & Ashton 1985, Cooper & Bra-
swell 1995, Cooper & Cooper 1995). Since
that time, many additional specimens from
the Neuse and Cape Fear river basins have
either been collected or have been recog-
nized in prior collections. They belong to
an undescribed species of Cambarus that
seems to have its closest affinities with con-
geners that occur in the Tennessee and Ohio
river drainages.
Abbreviations used in the text are as fol-
lows: j, juvenile; NC, North Carolina State
highway; NCSM, North Carolina State Mu-
seum of Natural Sciences, Raleigh; PCL,
postorbital carapace length; R, river; SR,
State secondary (county) road; TCL, total
carapace length; US, United States high-
way; USGS, United States Geological Sur-
vey; and UTM, Universal Transverse Mer-
cator coordinates.
Cambarus (Cambarus) davidi,
new species
Fig. 1, Table 1
Diagnosis.—Body and eyes pigmented,
eye small (X adult diam 1.7 mm, n = 30).
Carapace vaulted, thoracic section averag-
ing 1.3 times wider than deep (n = 52).
Rostrum acarinate; margins elevated, sub-
parallel, caudally thickened, strongly to
moderately constricted at base of acumen,
lacking marginal spines or tubercles; floor
(dorsal surface) of rostrum deeply concave,
ladlelike; acumen 24.5 to 49.1% (X =
33.6%, n = 52) length of rostrum, latter
13.0 to 19.3% GG =.16.2% nn: = 52): of
TCL. Areola 5.2 to 14.5 (X = 8.1, n = 76)
times longer than wide, constituting 35.4 to
432
41.8% (X = 37.6%, n = 76) of TCL and
42.2 to 47.3% (K = 44.1%, n = 45) of
PCL; areola sparsely punctate, with 2 (n =
22) to 3 (n = 64) punctations across nar-
rowest part. Cervical spines reduced to
multiple tubercles. Branchiostegal spine re-
duced to small tubercle; hepatic and sur-
rounding regions of carapace crowded with
tubercles. Suborbital angle obtuse to nearly
obsolete, almost always bearing small tu-
bercle; postorbital ridge short, cephalic
margin rounded and usually devoid of tu-
bercle. Antennal scale 2.0 to 3.6 (X = 2.5,
n = 50) times as long as broad, widest just
distal to midlength, lateral margin thick-
ened and with long distal spine.
Palm of chela of cheliped 1.5 to 1.8 (X
= 1.6, n = 51) times wider than deep, width
1.3 to 1.7 = 1.5, 2 = 51) tames length
of mesial margin; dorsolateral margin cos-
tate distally, without impression; mesial
margin of palm with 2, rarely 3, rows of
tubercles: mesial row of 6 to 8 (usually 7)
large, generally adpressed tubercles, sub-
tended dorsally by row of 1 to 5 (usually 4
or 5) smaller tubercles. Fixed finger of che-
la costate laterally, with well defined lon-
gitudinal ridges dorsally and ventrally; op-
posable surface of finger with row of 4 to
11 (usually 5 or 6) tubercles in addition to
subconical tubercle; dactyl 1.7 to 2.6 (X =
2.0, n = 51) times as long as mesial margin
of palm, with strong longitudinal ridge dor-
sally, weaker ridge ventrally; mesial margin
with prominent tubercles; opposable sur-
face with row of 6 to 14 (usually 7 to 9)
tubercles. Carpus of cheliped generally
lacking dorsomesial tubercles; merus with
prominent multiple dorsodistal tubercles
and often row of small squamous tubercles
along dorsal ridge.
Hook on ischium of third pereiopod of
males, that of form I male (Fig. 1K) uni-
ramous, overreaching basioischial articula-
tion and opposed by tubercle on basis; coxa
of fourth pereiopod of males with vertically
disposed, caudomesial boss.
In situ gonopods (Fig. 1G) symmetrical,
with abutted or slightly separated, tubercle-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
like proximomesial apophyses; proximola-
teral portion of gonopod set off from rest
of shaft by weak groove; in lateral aspect
(Fig. 1B), central projection curved over
90° to plane of shaft, untapered, with prox-
imally directed subapical notch; mesial pro-
cess inflated, symmetrically tapered, slight-
ly constricted at base of caudal third, with
subacute, caudally directed apex extending
slightly farther caudally than apex of cen-
tral projection; caudal process reduced to
swelling at caudodistal margin of shaft; in
mesial aspect (Fig. 1C), distal portion of
gonopod with essentially flat surface, few
setae at midlength.
Annulus ventralis of allotypic female
(Fig. 1H) 1.3 times broader than long, gen-
erally symmetrical and subrhomboidal; ce-
phalic margin convex and fused to sternum,
caudal margin subangular, free and capable
of slight movement; cephalic half of annu-
lus depressed, sloping, with narrow median
trough, flanked each side by low, poorly de-
fined ridge; ridges diverging caudally, si-
nistral ridge continuing obliquely and ter-
minating before joining caudosinistral wall,
dextral ridge curving caudodextrally to
merge with upper arm of heavy, C-shaped
caudodextral wall; transverse tongue prom-
inent, originating from sinistral wall near
caudal midline, continuing cephalically and
slightly obliquely before turning dextrally
and plunging into fossa beneath dextral
wall; sinus dissecting caudal wall near mid-
line.
Measurements of type specimens provid-
ed in Table 1.
Description of holotypic male, form I._—
Body and eyes pigmented, eye 1.8 mm
diam. Cephalothorax (Fig. 1A, D) subcy-
lindrical, thoracic section 1.2 times wider
than deep. Areola 9.2 times longer than
wide, constituting 39.3% of TCL (45.1% of
PCL), sparsely punctate, with 3 punctations
across narrowest part. Rostrum acarinate,
with slightly thickened caudal margins
moderately converging to base of acumen,
where moderately constricted; margins of
acumen slightly concave and converging to
VOLUME 113, NUMBER 2
Fig. 1. Cambarus (Cambarus) davidi, new species (all from holotypic male, form I, except E, EK from
morphotypic male, form II, and H, from allotypic female): A, lateral aspect of carapace; B, E, lateral aspect of
gonopod (first pleopod); C, EK mesial aspect of gonopod; D, dorsal aspect of carapace; G, caudal aspect of in
situ gonopods; H, annulus ventralis and postannular sclerite; I, dorsal aspect of distal podomeres of right cheliped;
J, antennal scale; K, hook on ischium of third pereiopod; L, epistome.
dorsally directed apical tubercle, which
reaching just beyond midlength of penulti-
mate podomere of antennular peduncle;
acumen comprising 30.2% of rostrum
length, latter constituting 14.2% of TCL;
floor of rostrum excavate, ladlelike, mod-
erately punctate, and ascending caudally
into broad dorsomedian depression of car-
apace; subrostral ridge strong, visible to
base of acumen in dorsal aspect.
434
Table 1.—Measurements (mm) of types of Cam-
barus (Cambarus) davidi, new species.
Holotyp- Morpho-
ic Allotypic typic
male female male
Carapace
Total length 30:3. 36:5: 4.3077
Postorbital length 26:4. 31.6 . 26.6
Length cephalic section 18.4 22.4 19.0
Width 19.7 "18:4 15S
Depth I2AGsl 1529 122
Length rostrum 4.3 5.6 4.9
Length acumen ee) 22 i
Length areola LO 14k ey,
Width areola 13 jai ino
Antennal scale
Length 4.5 5.4 Dye
Width 1.8 Jecdl 1.8
Abdomen
Length 29:6, 37.1 \B50:6
Width 1326 G4. -g122
Cheliped
Length lateral margin chela 24AA 26.7 23:8
Length mesial margin palm ies 8.8 fis
Width palm A i ha ad bl |
Depth palm 6.8 7.8 6.7
Length dactyl 1555 16:3); .452
Length carpus LO:0%, 1c) ee)
Width carpus iy? 8.0 6.7
Length dorsal margin merus r2:0) 13:87 212.2
Depth merus afa 8.5 6.9
Gondopod length 8.4 N/A 8.0
Postorbital ridge strong, groove essen-
tially lateral, cephalic margin with vestigial
tubercle. Branchiostegal spine reduced to
tubercle; suborbital angle obtuse, without
tubercle. Thoracic section of carapace dor-
sally and dorsolaterally punctate, laterally
with large, scattered granules; cephalic sec-
tion 1.5 times longer than areola and con-
stituting 60.7% of TCL, laterally crowded
with large tubercles and with row of small
tubercles along ventral margin of cephalic
section of cervical groove; gastric region
mostly glabrous. Cervical spine region on
right with 4 large and 3 smaller tubercles
(2 large and 2 smaller on left). Abdomen
slightly narrower and shorter than cepha-
lothorax. Proximal podomere of uropod
without spine or tubercle on lateral lobe,
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
with prominent caudomedian spine on me-
sial lobe; mesial ramus of uropod with me-
dian keel bearing strong caudal spine, tip of
which reaching caudal margin of ramus,
caudolateral margin with small spine; lat-
eral ramus with submedian keel bearing ter-
minal spine at transverse flexure, latter with
total of 17 fixed spines along margin and 1
long, movable sublateral spine. Telson with
1 long stationary and 1 long articulated
spine in each caudolateral corner of ce-
phalic section; caudal margin domelike.
Epistome (Fig. 1L) with symmetrical,
subtriangular cephalic lobe bearing short
cephalomedian projection; margins of lobe
uninterrupted, moderately thickened, lateral
apices thicker than rest; floor (ventral sur-
face) of lobe punctate and slightly convex,
concave along lateral margins; transverse
basal sulcus complete; central depression of
body broad, moderately deep, with cephal-
omedian fovea; lamellae punctate, lateral
margins subtruncate, with 2 large subacute
tubercles and 1 small rounded tubercle in
right caudolateral corner (1 large subacute
and 1 small rounded tubercle on left); zy-
goma moderately arched, pits elongate. An-
tennal peduncle with small cephalolateral
tubercle on basis, small ventral tubercle on
ischium; antennular peduncle with small,
laterally displaced subdistal spine on ven-
tral surface of basal podomere. Antennal
scale (Fig. 1J) 2.5 times longer than wide,
broadest just distal to midlength; lateral
margin thickened, terminating in long distal
spine, tip of which reaching distal margin
of penultimate podomere of antennular pe-
duncle; lamella ca. 1.3 times as wide as
thickened lateral margin; distal margin of
lamella subtransverse for most of width,
then sloping to widest point; mesial margin
subparallel to lateral margin for most of
length.
Third maxilliped with tip of endopodite
reaching about midlength of penultimate
podomere of antennal peduncle; exopodite
hirsute, tip reaching base of distal two-
thirds of merus of endopodite; cephalola-
teral corner of ischium slightly produced,
VOLUME 113, NUMBER 2
not spinelike; ventrolateral ridge flanked
mesially by row of punctations bearing
moderately long setae; lateral half of ischi-
um with punctations bearing short setae,
punctations most abundant on proximal
third; mesial half with long stiff setae large-
ly obscuring mesial margin, latter with 27
denticles on right. Right mandible with in-
cisor ridge bearing 7 denticles (6 on left).
Right chela (Fig. 11) 2.1 times longer
than wide; palm 1.7 times broader than
deep, width 1.6 times length of mesial mar-
gin; latter 29.9% of chela length, 47.1% of
dactyl length. Dorsal surface of palm punc-
tate; distolateral margin of palm and lateral
margin of fixed finger costate, area at junc-
ture of palm and finger with aggregation of
large punctations creating slight impres-
sion; lateral margin of palm rounded and
with row of large punctations. Ventral sur-
face of palm less punctate than dorsal, dis-
tolateral area with moderate depression and
aggregation of large punctations; lateral
eminence of articular ridge with distal tu-
bercle, none proximal to ridge. Mesial mar-
gin of right palm with mesial row of 7 ad-
pressed tubercles, proximal 3 of which with
elevated distal margins (same on left), me-
sial row subtended dorsally by row of 4
smaller tubercles (5 on left) and ventrally
by 1 small, squamous distal tubercle (3 on
left).
Fingers gaping in proximal three-fourths
of length, greatest width of gape about four-
fifths width of base of fixed finger; oppos-
able base of fixed finger with tuft of short
setae; both fingers slightly curved distov-
entrally in lateral aspect, dactyl very slight-
ly bowed in dorsal aspect. Mesial margin of
right dactyl bearing row of 4 prominent and
4 weaker tubercles on proximal half, distal
half of margin punctate; dorsal surface of
dactyl with low, rounded longitudinal ridge,
flanked mesially by punctate groove, later-
ally by row of large, spaced punctations;
ventral surface with similar longitudinal
ridge; opposable surface with 8 tubercles,
fourth from base very large and slightly dis-
placed ventrally (7 tubercles on left, third
435
from base largest); denticles in 2 or 3 rows
from near tip of finger to sixth tubercle
from base, single row from there. Fixed fin-
ger with very strong dorsomedian ridge
flanked mesially by deep punctate groove
and second narrower ridge, laterally by
deep punctate groove; lateral margin with
irregular row of large punctations; ventral
surface with very strong longitudinal ridge,
flanked both sides by row of large puncta-
tions; opposable margin with subconical tu-
bercle ventral to denticles at base of distal
third of finger, and 5 tubercles proximal to
subconical one, third from base more prom-
inent than others (4 tubercles on left finger,
third from base very large); denticles in 2
or 3 rows from tip of finger to subconical
tubercle, single row from there.
Carpus (Fig. 11) 1.4 times as long as
wide, 1.4 times as long as mesial margin of
palm; carpus dorsally with long, deep,
slightly oblique sulcus, lateral and mesial to
which surface punctate; mesial margin with
large distal spine and prominent proximal
tubercle; ventral surface with stout, rounded
tubercle at lateral articular condyle, similar
distomedian tubercle, and 1 small tubercle
proximomesial to latter. Right merus 1.7
times longer than deep; dorsal surface with
3 prominent and 2 smaller subdistal tuber-
cles (2 prominent and 1 smaller on left),
and small squamous tubercles along much
of dorsomedian ridge; ventrolateral ridge
with 2 small acute tubercles and 1 minus-
cule tubercle near articular condyle (1 small
acute tubercle and minuscule distal tubercle
on left); ventromesial ridge with 10 spini-
form tubercles, distalmost one somewhat
larger than others; ischium with 3 small
ventral tubercles (4 on left).
Palm and fingers of chela of second pe-
reiopod hirsute. Ventral margins of pleura
subtruncate or slightly rounded, caudoven-
tral corners slightly angular, caudal margins
rounded; terga very punctate, except artic-
ular cephalic portions glabrous. Sternites
between third and fourth coxae with very
dense, matted setae, covering distal ends of
in situ gonopods.
436
Gonopods (Fig. 1B, C, G) as described
in “‘Diagnosis.’’ Length of gonopod 27.7%
of: TCL: G8 of PCE):
Description of allotypic female.—Except
for secondary sexual characters, differing
from holotypic male in following respects:
Areola 6.7 times wider than long, consti-
tuting 38.6% of TCL (44.6% of PCL). Acu-
men comprising 39.3% of rostrum length,
latter constituting 15.3% of TCL. Suborbit-
al angle nearly obsolete, with small tuber-
cle. Cervical spine region on both sides of
carapace with 3 prominent tubercles, largest
of those on right side subacute. Caudolater-
al corner of cephalic section of telson with
2 spines on right, 3 on left. Cephalic lobe
of epistome subcordiform; lamellae with
single tubercle each caudolateral corner; zy-
goma strongly arched. Antennal peduncle
lacking tubercle on basis; antennal scale 2.6
times longer than wide, lamella about 1.5
times as wide as thickened lateral margin.
Incisor ridge of right mandible bearing 8
denticles. Right chela 2.0 times longer than
wide, palm length 33.0% of chela length,
54.0% of dactyl length. Opposable surface
of right dactyl with 10 tubercles (9 on left);
opposable surface of both fixed fingers with
6 tubercles in addition to subconical one,
fourth from base largest. Carpus of cheliped
1.3 times as long as mesial margin of palm;
merus 1.6 times longer than deep, dorsal
surface with 2 prominent and 2 smaller sub-
distal tubercles (same on left); ventrolateral
ridge with 5 small tubercles (same on left),
ventromesial ridge with 8 spiniform tuber-
cles (9 on left).
Annulus ventralis (Fig. 1H) as described
in “‘Diagnosis.”’ In addition, first pleopods
short, reaching caudal margin of annulus
when abdomen flexed; annulus about 3
times wider than postannular sclerite, which
elongate, ventrally domed, punctate.
Description of morphotypic male, form
II.—Differing from holotypic male in fol-
lowing respects: Thoracic section of cara-
pace 1.3 times wider than deep. Areola 6.2
times as long as wide, constituting 38.1%
of TCL (44.0% of PCL). Margins of ros-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
trum less constricted at base of acumen,
converging at ca. 45°, acumen comprising
24.5% of rostrum length, latter constituting
16.0% of TCL. Suborbital angle obtuse and
with very small tubercle. Cephalic section
of carapace 1.6 times longer than areola and
constituting 61.9% of TCL. Cervical region
on both sides of carapace with 3 prominent
and 1 moderate tubercle, ventralmost of
which on right side subacute. Transverse
flexure of lateral ramus of right uropod with
16 spines in addition to sublateral one. Lat-
eral corners of lamellae of epistome with 1
moderate tubercle on right, 2 on left; zy-
goma strongly arched. Antennal scale 2.9
times wider than long, lamella ca. 1.5 times
width of thickened lateral margin, distal
margin moderately declivous. Length of
palm of cheliped 31.5% of chela length,
49.3% of dactyl length. Mesial margin of
palm with mesial row of 6 adpressed tu-
bercles, subtended dorsally on right palm
by row of 3 moderate tubercles (4 on left)
and ventrally by 4 small squamous tuber-
cles (2 on left). Opposable surface of dactyl
with 9 tubercles on right, 8 on left, basal-
most largest. Opposable surface of fixed
finger with 6 tubercles on right in addition
to subconical one, 7 on left, third from base
largest. Carpus 1.3 times as long as palm.
Merus 1.8 times longer than deep; dorsal
surface of right merus with patch of 5
prominent subdistal tubercles and 5 smaller
tubercles proximal to them (2 moderate and
6 smaller on left); ventrolateral ridge on
right with 2 subacute and 2 other tubercles
(2 small, and 1 minuscule articular tubercle,
on left); ventromesial ridge on right with 10
small tubercles (9 on left).
Hook on ischium of third pereiopod
weak, not overreaching basioischial articu-
lation, opposed by tubercle on basis; boss
on coxa of fourth pereiopod moderately de-
veloped.
Gonopod (Fig. 1E, F) length 26.1% of
TCL. In situ gonopods with weak, separat-
ed proximomesial apophyses; mesial pro-
cess noncorneous, bulbous, distal surface
creased near extruded tip; in lateral aspect,
VOLUME 113, NUMBER 2
gonopod with weak juvenile suture; central
projection noncorneous, curved 90° to
shaft, tapered to subacute tip; mesial pro-
cess tapered, triangular in outline, tip di-
rected caudally and inclined slightly disto-
laterally. Setae on sternites between third
and fourth coxae not dense.
Color notes.—Adult ground color usu-
ally dark olivaceous, sometimes light
brown with orange or tan overtones. Ce-
phalic section of carapace often lighter in
color than thoracic section. Hepatic region
with orangish midlateral streak just cephalic
to cervical groove. Margins of rostrum out-
lined in tan; antennal scale pale orangish-
tan with dark lateral margin, antennal fla-
gellae green. Most tubercles, spines, and
granules of carapace and chelipeds tan to
orangish. Dorsal surface of cheliped green-
ish or olivaceous; articular ridges of chela
pinkish or orangish, ventral surface of chela
pale grayish-tan. Lateral surface of entire
propodus (palm and fixed finger) strikingly
colored, varying from pinkish-tan to
creamy orange or yellow. Tips of fingers of
chelipeds pale orange or orangish-tan, color
not subtended by black band. Proximal po-
domeres of other pereiopods pale tan to
light brown with darker mottling, distal po-
domeres greenish or bluish, fingers of che-
lae of second and third pereiopods pale
blue. Cephaloventral structures bluish-gray,
except epistomal zygoma almost white. An-
nulus ventralis usually pale, mottled with
orange in one female.
Cephalicmost tergite of abdomen with
transverse dark brown or black rectangular
band. Lighter colored adults and all juve-
niles with dark, diagonal blotch each side
of caudal margin of thoracic section of car-
apace, blotches extending onto dorsolateral
surfaces of two adjacent tergites as short,
curved markings. Series of short, dark bars
producing interrupted dorsolateral stripe
each side of abdomen. Other dorsal surfaces
of abdomen with scattered dark spotting.
Ventrolateral pleura of abdomen with pale
pink cephalic area and narrow oblique or
V-shaped black bar dorsal to it, series of
437
these bars producing zigzag lateral stripe
each side of abdomen. Juveniles generally
paler than adults, color patterns in most re-
spects similar but more vivid.
Type _ locality.—North Carolina, Wake
County, small intermittent stream entering
cove along western shore of Falls Lake (im-
poundment of Neuse River), ca. 1.4 air km
NW of western end of NC 98 bridge & ca.
2.6 air km W of Stony Hill (Bayleaf 7.5’
USGS quadrangle, UTM zone 17, 3984850/
712190).
The shallow stream, which lies at the
bottom of a steep ravine in a hardwood for-
est, is seasonally intermittent and has a
maximum width of about 1.2 m. All spec-
imens from this locality were found in the
mud of shallow residual pools under large
rocks when water levels were very low.
Disposition of types.—The holotypic
male, allotypic female, and morphotypic
male are in the crustacean collection of the
NCSM (catalogue numbers NCSM C-4413,
C-4414, and C-2656, respectively), as are
paratypes consisting of 2 6 I, 15 ¢ II, 6 j
S18 iS and 57 <2.
Range and specimens examined.—Ap-
parently limited to the upper Neuse and
Cape Fear river basins in the eastern Pied-
mont Plateau of North Carolina. Voucher
specimens (n = 107), all in the crustacean
collection at NCSM (catalogue numbers in
parentheses), have been collected at the fol-
lowing localities.
Neuse River Basin: Durham Co.; upper
trib Little R near Durham, 1 ¢ II (C-4742),
13 Feb 1995, coll. T. Cuffney. Orange Co.;
small intermittent stream in headwaters
West Fork Eno R at SR 1358, 2.4 air km E
of Cam, 3:7 2i(C-3425), 25. Jal! 1995,. coll.
M.A. Hartman, M.E. Savacool. Wake Co.;
seep entering small trib New Light Crk
along SR 1918, 0.3 km SW of jct SR 1909,
ca. 7.4 air km NNE of Bayleaf, 2 jd (C-
44), 18 Feb 1976, coll. A.L. Braswell
(ALB), N. Murdock; type locality, 1 ¢ I
(C-2656), 1 jd (C-2657), 4 Jul 1994, coll.
Eee Copper (DGC), JEC, Ligd 5.3. 2)(C-
2779), 1 j2 (C-2780), 3 jd (C-2781), 18
438
Aug 1993, coll. DGC, JEC, 1 6 I (C-4413),
8. dT 6 G56 “Py Tgln(GE-3795)) 14: Jun
1997, coll. DGC, 1 2 (C-4553), 20 Jun
1998, coll. DGC; spring on small trib Low-
er Barton Crk between SR 1005 & SR
1844, SW of Bayleaf, 2 32 (C-3055), 6 Apr
1996;scollS: YirkayDG@asjGlk 2a 248
(C-3293), 4 Jul 1996, coll. D. DeOliveira
(DD), DGC; spring on S shore Lower Bar-
ton Crk, W of SR 1005, ca. 1.6 km NNW
of ‘center of Bayleaf, 1) d7ie(E-3333), bh!
Aug 1996, coll. DGC; small spring entering
Falls Lake, ca. 0.8 km N of entrance of
Lower Barton Crk into lake, 2 ¢ II, 1 2,3
j2 (E-5IS Doo twApr: 1999:.-coll...DGE:
“Raleigh,” 2 2 (C-3143), 28 Nov 1924,
coll. C.S. Brimley, W.B. Mabee; small in-
termittent stream entering lake at Schenck
Forest, Raleigh, 1 ¢ II (C-3471), 25 Oct
1996, coll. DGC, 1 2 (C-3603), 4 Apr
1997, coll. DGC; small trib to Richland Crk
near SW side of Reedy Creek Rd, Schenck
Forest, Raleigh, 2 2 (C-5077), 10 Apr
1999, coll. DGC; small stream E of Jeffrey
Dr off Lake Wheeler Rd, SE of Lake
Wheeler Raleich, 3.9. Sao 4-371 7).20
May 1997;:colls DD; DGE;, Ind. 11.546, 4
j2 (C-3766), 1 2 (C-4414), 24 May 1997,
coll. DGC; Crabtree Crk below Duraleigh
Rd, Raleigh, 1 j2 (C-4591), 30 Aug 1998,
coll. D.A. Jackan.
Cape Fear River Basin: Alamance Co.;
spring on trib Toms Crk, Scott farm off SR
1612, 0.8 km NW of Union Ridge, 1 6 I,
1 2 in amplexus (C-45), 2 Mar 1976, coll.
ED. Scott (FDS); burrow at 403 Glen Ra-
ven Rd, Burlington, 1 6 II (C-3618), 25 Jun
1993, coll. C. McGrath (CM). Caswell Co.;
Benton Branch between SR 1103 and 1105,
S of SR 1100, ca. 1.9 air km SW of town
of, Stoneycréek; 4)621Ri ij Gein (C789);
14 May 1975, coll. FDS, JEC. Chatham
Co.; small, intermittent upper trib New
Hope Crk at SR 1716, 6.6 km NNE of jet
US 64, ca. 10.4 air km ENE of center of
Bynum, | jd (C-3026), 1 Apr 1986, coll.
D.R. Lenat (DRL), T. MacPherson, 1 j 2 (C-
3106),,5.-Oct,.1992,.colk \E Miss 7ou (Ez
3748) U. Feb41993 calls -CMs kid he (C-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
4464), 13 Feb 1998, coll. DRL, D. Penrose
(DP); upper trib Robeson Crk at US 64,
Pittsboro, 1 2 (C-4316), 6 Mar 1997, coll.
DP. Harnett Co.; upper trib Kenneth Crk at
SR 1447, NE of Rawls, 2 6 II, 1 jd (C-
2963), 28 Aug 1991, coll. N. Medlin (NM),
DP. Rockingham Co.; spring on small trib
Benaja Crk, ca. 0.3 air km ESE of jct SR
2426 and 242734: Cp iniiss 2) 24252 €
3104), 1 3 If (C-3105), 3 Jun 1976, coll.
M.R. Cooper, JEC; drainage ditch in flood-
plain Haw R, Camp Guilrock, ca. 19.2 air
km SE of Monroeton, 1 2 (C-4843), Mar
1998, coll. A.B. Somers & students; Little
Troublesome Crk at SR 2600, ca. 1.1 air km
W of Williamsburg, 1 2 (C-4912), 6 Apr
1998, coll. B. Tracy, NM, L. Eaton, DP.
Variations.—In addition to those ad-
dressed in the “Diagnosis,”’ the following
variations have been recorded. The margins
of the rostrum are usually abruptly or at
least moderately constricted at the base of
the acumen, but in seven specimens the
margins, while increasing in convergence,
are not notably constricted. The number of
prominent cervical tubercles varies from
one to six (usually three to five), and in
some specimens at least one of these tuber-
cles is spiniform. In three individuals, the
usual small tubercle on the suborbital angle
is absent, and in several the angle itself is
subacute. Nearly all specimens have a mi-
nuscule tubercle or very weak spine on both
the basis and ischium of the antennal pe-
duncle, but five lack a tubercle on the is-
chium and two lack a tubercle on the basis.
The width of the lamella of the antennal
scale ranges from approximately 1.1 to 2.0
(X = 1.4) times the width of the thickened
lateral margin. The distal margin of the la-
mella is usually either subtransverse or
moderately sloping for much of its width,
but in three females it is strongly declivous
from the base of the distolateral spine to the
mesial margin.
Most individuals have two spines in each
caudolateral corner of the cephalic section
of the telson, but seven of them have two
spines in one corner and three in the other,
VOLUME 113, NUMBER 2
and two have two spines in one corner and
one in the other. The lateral lobe of the
proximal podomere of the uropod normally
lacks a spine or tubercle, but 12 specimens
have a very small spine or acute tubercle
on this lobe. The spine on the mesial lobe
of this podomere varies in size from very
small to moderate, and is absent in one an-
imal.
The chela of form I males is longer than
that of mature form II males and females,
averaging 81% of TCL. In form II males
the average is 71.6% and in females it is
70.8%. The largest tubercle on the oppos-
able surface of the fixed finger varies from
the third to the fifth from the base, but in
most specimens the fourth tubercle is much
larger than the others. The largest tubercle
on the comparable surface of the dactyl
varies from the first to the fifth, but in most
it is the fourth, and the largest tubercle is
almost always offset toward the ventral sur-
face. The dorsal surface of the merus bears
from one to eight prominent subdistal tu-
bercles, with the usual number being three
or four. Most specimens have from two to
four weaker tubercles just proximal to the
more prominent ones, and many have squa-
mous to subsquamous tubercles along at
least part of the dorsal ridge. The number
of tubercles on the ventrolateral ridge of the
merus ranges from two to five (usually
three), and the distalmost is very small or
vestigial. The number of tubercles on the
ventromesial ridge ranges from six to elev-
en (usually nine or ten), and the distalmost
is seldom much larger than the largest of
the others.
In 17 adult females the width of the an-
nulus ranges from 1.2 to 1.8 (X = 1.5)
times its length, and the cephalomedian
trough varies from moderately wide to
nearly obliterated. The thick, C-shaped wall
of the annulus, beneath which lies the deep-
est part of the fossa, is dextral in 22 fe-
males, sinistral in eight others.
The floor (dorsal surface) of the rostrum
of juveniles and some subadults, while no-
tably excavate, is often less ladlelike than
439
it is in larger, mature animals. Also in these
smaller individuals, the setae on the oppos-
able surface of the fixed finger of the che-
liped are far longer and more dense than
they are in adults, often filling the space
between the fingers and obscuring the tu-
bercles on both.
Size.—The largest specimens collected
are two females measuring 50.7 and 50.0
mm TCL (44.4 and 44.5 mm PCL), both
from the Haw River subdrainage of the
Cape Fear River basin. The next largest
specimen is a form I male, which measures
42.5 mm TCL (37.3 mm PCL). The largest
form II male measures 33.4 mm TCL (28.7
mm PCL).
Life history notes.—A form I male was
collected at the type locality on 14 June
1997, one was taken in Chatham County on
13 February 1998, and one was found in
amplexus with a female in Alamance Coun-
ty on 2 March 1976. No females with at-
tached ova or young have yet been seen,
but one measuring 32.5 mm TCL, taken on
4 April 1997, had all cement glands highly
developed.
Crayfish associates.—Seldom have other
crayfishes been found in the same habitats
with C. (C.) davidi. At a few localities,
however, a number of specimens of Cam-
barus (Depressicambarus) latimanus (Le-
Conte, 1856), and of at least one of the spe-
cies in the complex subsumed under Cam-
barus (Puncticambarus) acuminatus Faxon,
1884, have been found. At two sites, juve-
nile C. (D.) reduncus were collected. While
juvenile Cambarus (Lacunicambarus) di-
ogenes Girard, 1852, have been taken from
under cover near the mouth of the stream
at the type locality, and chimneyed burrows
of this species may be seasonally abundant
along the nearby shoreline of Falls Lake,
this burrowing species has not been found
with C. (C.) davidi.
Relationships.—Ilt appears to me that C.
(C.) davidi has its strongest affinities with
Cambarus (Cambarus) bartonii (Fabricius,
1798). Students of American crayfishes,
however, have long been cognizant of the
440
taxonomic perplexities presented by the
broadly distributed and highly variable pop-
ulations currently assigned to this species.
Two subspecies have been described, Cam-
barus (Cambarus) bartonii cavatus Hay,
1902, and Cambarus (Cambarus) bartonii
carinirostris Hay, 1914, but opinions anent
their validity have for years fluctuated (see
Faxon 1914, Ortmann 1931, Hobbs 1972,
1989, Bouchard 1976, Thoma & Jezerinac
1982, Jezerinac 1985, Fitzpatrick 1983, Jez-
erinac & Thoma 1984, Jezerinac et al.
1995, Cooper 2000). The status of C. (C.)
b. cavatus remains controversial, but C.
(C.) b. carinirostris has been elevated to
species status (Thoma & Jezerinac 1999).
Although it is still difficult at this time to
establish precise diagnostic parameters for
these taxa throughout their ranges (what-
ever those ranges might be), current diag-
noses must be used in assessing the rela-
tionships of C. (C.) davidi.
In his brief description of C. b. cavatus,
whose type locality is the Powell River
(Tennessee-Ohio river drainage) near Ta-
zewell, Claiborne County, Tennessee, Hay
(1902:435) emphasized its “‘deeply exca-
vated rostrum,’’ an areola that is “‘narrower
and more thickly punctate than in C. bar-
toni bartoni,”’ and a carapace that is “‘more
nearly cylindrical.’’ Except for the “‘more
thickly punctate’’ areola, a number of the
characters displayed by C. (C.) davidi in-
dicate a possible relationship with the pro-
genitors of “‘topotypic”’ C. (C.) b. cavatus,
although their similarities could just as well
be a result of convergence. In Ohio and
West Virginia, this putative subspecies ap-
parently lacks the deeply excavate rostrum,
and displays other characters that differ
from those of the “‘typical’’ form (Jezerinac
1985, Jezerinac et al. 1995).
The combination of a narrow (but not
obliterated), sparsely punctate areola, a la-
dlelike rostrum, and a vaulted, unflattened
carapace will serve to separate C. (C.) dav-
idi from all other members of the subgenus
except some C. (C.) b. cavatus, and some
upland southern populations currently as-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
signed to C. (C.) b. bartonii (s.1.). Camba-
rus (C.) davidi differs from C. (C.) b. ca-
vatus and most populations of C. (C.) b.
bartonii (s.1.) in its obtuse to nearly obso-
lete suborbital angle, the shape of its anten-
nal scale, and the presence of multiple tu-
bercles on the dorsal surface of the merus
of the cheliped. In some montane popula-
tions of C. (C.) b. bartonii (s.1.) the cara-
pace is relatively vaulted and the rostrum is
deeply excavate and somewhat ladlelike.
These populations, however, characteristi-
cally have areolae that are broader and have
more punctations across the narrowest part
than does the areola of C. (C.) davidi. Also,
in nearly all C. (C.) bartonii (s.1.) the la-
mella of the antennal scale is much narrow-
er than it is in C. (C.) davidi, and its distal
margin is usually quite declivous from the
base of the lateral spine to the mesial mar-
gin. In addition, C. (C.) bartonii (s.1.) sel-
dom displays prominent multiple tubercles
on the dorsal surface of the merus.
Roger E Thoma, whose knowledge of
subgenus Cambarus is extensive, has sug-
gested that a comparison of C. (C.) davidi
with the burrower, Cambarus (Cambarus)
ortmanni Williamson, 1907, would be use-
ful. That species differs from C. (C.) davidi
in many ways, including the following:
Areola of C. (C.) ortmanni generally oblit-
erated or nearly so, constituting 41.0 to
44.0% of TCL; suborbital angle obsolete
and lacking tubercle; cephalothorax mark-
edly longer than abdomen; subdistal spine
on mesial margin of carpus of cheliped
thick and knoblike rather than long and
acute; cervical region of carapace lacking
strong, multiple tubercles; ventrolateral
ridge of merus of cheliped usually lacking
tubercles; and annulus ventralis and form I
male gonopod quite different in configura-
tion.
Remarks.—Current evidence indicates
that the range of C. (C.) davidi, limited to
parts of the eastern Piedmont Plateau in two
endemic North Carolina river basins, is dis-
junct from that of other members of the
subgenus. The nearest known North Caro-
VOLUME 113, NUMBER 2
lina populations of C. (C.) bartonii (s.1.) are
in the mountains and eastern foothills.
Whether or not C. (C.) davidi is indeed al-
lopatric, however, will only be revealed by
more extensive field work throughout the
Piedmont Plateau. Specimens of subgenus
Cambarus from seeps and burrows in the
Dan River subdrainage of the Roanoke Riv-
er basin resemble C. (C.) davidi in many
respects, differ in others. The sample size
of available adult specimens is far too small
for accurate assignment of the Roanoke ma-
terial at this time.
Cambarus (C.) davidi has yet to be found
in the Tar-Pamlico River basin, whose cray-
fish fauna is as well known as that of the
Neuse and is nearly identical (Cooper &
Braswell 1995). Unfortunately, almost no
sampling has been done in appropriate hab-
itats within the Tar-Pamlico basin.
Etymology.—I take great pleasure in
naming this species for my son, David
George Cooper, an avid naturalist who
brought the species to my attention and who
spent many hours tromping in mud and
muck to collect quite a few of the existing
specimens.
Suggested vernacular name: Carolina la-
dle crayfish.
Acknowledgments
I am indebted to David G. Cooper for his
enthusiastic and persistent field work, as
well as to those other biologists who col-
lected specimens (their names are provided
in the section on ‘‘Range and specimens ex-
amined’’). I also express my sincerest grat-
itude to Alvin L. Braswell, John E. Cooper,
Jr., Martha Riser Cooper, and Don Howard,
for their unstinting assistance. The manu-
Script was greatly improved by the reviews
of Roger E Thoma, Rafael Lemaitre, and
an anonymous referee. Nancy Childs,
NCSM, provided technical assistance in the
final preparation of Figure 1.
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zoogeography of Cambarus bartonii cariniros-
tris Hay, 1914 (Crustacea: Decapoda: Cambar-
idae).—Proceedings of the Biological Society
of Washington 112:97—105.
Williamson, E. B. 1907. Notes on the crayfish of Wells
County, Indiana, with a description of a new
species. Pp. 749-763 in 31st Annual Report of
the Department of Geology and Natural Re-
sources of Indiana, 1906.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):443—457. 2000.
Two new species of Hyalella (Crustacea: Amphipoda: Hyalellidae)
from Death Valley National Park, California, U.S.A.
Adam J. Baldinger, William D. Shepard, and Doug L. Threloff
(AJB) Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge,
Massachusetts 02138-2902, U.S.A.; (WDS) Department of Biological Sciences,
Sacramento State University, Sacramento,
California 95819-6077, U.S.A.; (DLT) United States Department of the Interior, National Park
Service, Death Valley National Park, Death Valley, California, 92328, U.S.A.
Abstract.—Two new species Hyalella (Hyalella) muerta and Hyalella (Hy-
alella) sandra are described from Death Valley National Park, California. Hy-
alella (H.) muerta, the first North American hypogean hyalellid, is blind, lacks
dorsal mucronations and antenna | is longer than antenna 2. Hyalella (H.)
sandra collected from nearby epigean waters, also lacks dorsal mucronations,
but has normal eye pigmentation and antenna 1 is shorter than antenna 2.
Populations of both species rarely, if ever, coexist together in Death Valley
National Park.
Five species in the genus Hyalella occur
in the continental United States. These are
Hyalella (Hyalella) azteca (Saussure,
1858); Hyalella (Hyalella) inermis Smith,
1874; Hyalella (Hyalella) montezuma Cole
& Watkins, 1977; Hyalella (Ayalella) tex-
ana Stevenson & Peden, 1973; Hyalella
(Hyalella) longicornis Bousfield, 1996. All
are epigean species, have normal eye pig-
mentation and have antenna 1 shorter than
antenna 2. Only H. (H.) inermis and H. (H.)
longicornis lack dorsal mucronations.
Examination of recently collected mate-
rial from Death Valley National Park, Cal-
ifornia clearly shows the presence of two
new species of Hyalella, both of which are
newly described in this paper.
In the figures, body parts are marked by
the following abbreviations: A, antenna;
Gn, gnathopod; LL, lower lip; UL, upper
lip; Md, mandible; Mx, maxilla; Mxpd,
maxilliped; P, pereopod; T, telson; U, uro-
pod; Pl, pleopod; R, right; L, left. Type ma-
terial is deposited in the Departments of In-
vertebrate Zoology at the following muse-
ums: National Museum of Natural History
(USNM); Museum of Comparative Zoolo-
gy, Harvard University (MCZ) and Califor-
nia Academy of Sciences (CASIZ).
Hyalella (AHyalella) muerta, new species
Figs. 1-5
Diagnosis.—Eyes absent. Antenna 1 lon-
ger than antenna 2. Accessory flagellum ab-
sent. Pereon lacking dorsal mucronations or
spines. Sternal gills reduced in size and pre-
sent on pereonites 3—7. Maxilla 1, inner
plate narrow with 2 terminal plumose setae.
Male gnathopod 2, carpus with strong pos-
terior lobe with 4 submarginal setae; pro-
podus robust and much larger than male
gnathopod 1, palm with rectangular tooth
near hinge of dactylus. Telson rounded,
with 4 distal marginal setae.
Description of male.—Body lacking dor-
sal mucronations or spines. Eyes absent.
Head cuboidal, equal to pereonites 1 and 2
in length. Coxae 1—4 enlarged, subquadrate
with distomarginal setae, coxae 5—6 with
distinct anterior and posterior lobes. Sternal
gills small, present on pereonites 3-7.
444
Antenna 1, 57% of total body length, pe-
duncular ratio 1:0.7:0.6, flagellum 9 artic-
ulate; accessory flagellum absent. Antenna
2 shorter than antenna 1, 49% of body
length, peduncle nearly equal in length to
head, articles 4 and 5 equal in length, fla-
gellum 8 articulate.
Upper lip rounded, anterior margin with
fine setae. Mandibles lacking palp; left in-
cisor and lacina with 4 teeth, molar normal.
Right incisor with 5 teeth, outer the largest
and bifid, lacina with 3 teeth, spine row
with 2 plumose setae, molar normal, with
accessory plumose seta. Lower lip large,
lacking inner lobes. Maxilla 1, inner plate
narrow with 2 terminal plumose setae, inner
margin finely setose; outer plate with 8
strong apical serrate spines; palp vestigial,
1 articulate. Maxilla 2, plates subequal in
width; inner plate with 1 marginal and 12
apical plumose setae, outer margin with fine
setae distally; outer plate with 10 apical
plumose setae, both margins with fine setae.
Maxilliped, inner plate with 3 strong apical
teeth, inner margin with 3 plumose setae
distally; outer plate with 7 apical plumose
setae, with marginal and submarginal plu-
mose setae; palp 4 articulate, nearly 3X
length of outer plate, inner margin of article
2 with distal plumose setae, article 3 with
distal setae, dactylus triangular with 3 distal
setae.
Gnathopod 1, basis elongate broadening
distally, posterior lobe of carpus with 6
marginal setae; propodus rectangular, wider
and longer than carpus; palm and hind mar-
gin equal in length, palm with marginal and
submarginal setae. Gnathopod 2, basis elon-
gate; posterior lobe of carpus strong with 4
submarginal setae; propodus robust and
much larger than that of gnathopod 1, distal
anterior margin with triangular projection;
palm with rectangular tooth near hinge of
dactylus, with marginal and submarginal
setae. Pereopods 3 and 4 similar in shape
and size; carpus of both with distal anterior
lobe and 2 setae. Pereopod 5, approximate-
ly 70% length of pereopod 6. Pereopod 7
slightly larger that pereopod 6. Pereopods
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
5—7 bases expanded posteriorly, stronger in
pereopod 7; carpus with distal posterior
lobe, with 3—4 spines.
Pleopods 1-3 long and slender, pedun-
cles with 2 coupling hooks, rami with plu-
mose setae. Uropod 1, peduncle outer mar-
gin with 4 bifurcate spines, inner margin
with distal marginal seta; inner ramus with
4 apical spines; outer ramus with 5 apical
and 2 marginal spines. Uropod 2, 60% the
length of uropod 1, peduncle with 2 mar-
ginal spines and 1 marginal seta; inner ra-
mus with 6 apical spines; outer ramus with
5 apical spines. Uropod 3, peduncle with 3
distal marginal spines; ramus with 2 apical
spines and 1 apical seta. Telson rounded
with 4 distal marginal setae.
Female.—All features same as male ex-
cept as noted below. Antenna 1, 43% of
total body length; flagellum 9 articulate.
Antenna 2, 34% of total body length; fla-
gellum 9 articulate. Gnathopod 2 and pe-
reopods 3—5 with oostegites. Gnathopod 1
resembling male, posterior lobe of carpus
with 8 marginal setae. Gnathopod 2, basis
elongate; posterior lobe of carpus with 6
submarginal setae; propodus longer than
wide, palm and hind margin distinct.
Etymology.—The specific epithet is de-
rived from the Spanish muerta as a noun in
apposition for the word death referring to
Death Valley.
Material examined.—Male holotype,
3.28 mm, USNM 230433, Texas Spring,
just uphill of the Texas Spring Camp-
ground, Death Valley National Park, Inyo
County, California, William D. Shepard, 20
Jul 1994. Female (ovigerous) allotype, 3.28
mm, USNM 230434, same data as holo-
type. Male paratype, 3.28 mm, USNM
230435, same data as holotype. Female
(ovigerous) paratype, 3.24 mm, USNM
230436, same data as holotype. 6 males, 8
females (4 ovigerous) paratypes, USNM
230437, same data as holotype. Male (4.00
mm) and female (3.60 mm), paratypes,
USNM 230438, Texas Spring tunnel, Death
Valley National Park, Inyo County, Cali-
fornia, 36°27'27.54"N, 116°50'14.44’W,
VOLUME 113, NUMBER 2
Perea ae
Es an Ae
Fig: 1.
Doug L. Threloff, 19 Dec 1997. Paratypes,
MCZ 25390, CASIZ 121603, Texas Spring
tunnel, Death Valley National Park, Inyo
County, California, 36°27'27.54’N,
116°50'14.44"W, D. L. Threloff, 7 Dec
if. Paratypes, MCZ 25391, CASIZ
121604, Texas Spring tunnel, at discharge
point, Death Valley National Park,
Inyo County, California, 36°27'26.18’N,
116°50'16.93"W, D. L, Threloff, 19 Dec
Tu9T:
Remarks.—Hyalella (H.) muerta is mor-
phologically similar to Hyalella (Mesohy-
alella) anophthalma Ruffo, 1957 and Hy-
alella (Mesohyalella?) caeca Pereira, 1989;
the former is known from a cave in Vene-
zuela and the latter from Brazil. Both spe-
cies have sternal gills on pereonites 2—7 and
antenna 2 is longer than antenna 1. In con-
trast, H. (H.) muerta has sternal gills on
pereonites 3—7 and antenna | is longer than
antenna 2.
445
Hyalella (Hyalella) muerta, male, 3.28 mm, USNM 230435.
Hyalella (Hyalella) sandra, new species
Figs. 6—10
Diagnosis.—Eyes present. Antenna |
Shorter than 2, flagellum 10-11 articulate;
accessory flagellum absent. Antenna 2
elongate, flagellum 20—24 articulate. Pere-
on lacking dorsal mucronations or spines.
Sternal gills on pereonites 3—7, approxi-
mately % the size of coxal gills. Maxilla 1,
inner plate narrow with 2 terminal plumose
setae. Male gnathopod 2, carpus with strong
posterior conical lobe with marginal setae.
Uropod 3, ramus with 7 distal spines. Tel-
son, rounded with 2 distal submarginal
spines.
Description of male.—Body lacking dor-
sal mucronations or spines. Eyes pigment-
ed. Head cuboidal, subequal to pereonites |
and 2 in length. Coxae 1-3 enlarged, quad-
rate with distomarginal setae, coxae 5—6
with distinct anterior and posterior lobes.
446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
sal PP
Rigs.
Hyalella (Hyalella) muerta, male, 3.28 mm, USNM 230435.
VOLUME 113, NUMBER 2
Sternal gills on pereonites 3—7, approxi-
mately % the size of coxal gills.
Antenna 1, 25% of total body length, pe-
duncular ratio 1:1:0.8, flagellum 11 articu-
late each with small aesthetasc-like spines;
accessory flagellum absent. Antenna 2
much longer than antenna 1, 50% of body
length, peduncle articles 3 and 4 equal in
length to 5, flagellum 20 articulate.
Upper lip rounded, anterior margin with
fine setae. Mandibles lacking palp, molars
normal with accessory plumose seta; left in-
cisor and lacina with 5 teeth, spine row with
3 plumose setae. Right incisor with 5 teeth,
lacina with 3 teeth. Lower lip large, setose
and lacking inner lobes. Maxilla 1, inner
plate narrow, with 2 terminal plumose se-
tae; outer plate with 9 strong apical serrate
comb-like spines; palp vestigial, 1 articu-
late. Maxilla 2, plates subequal in width;
inner plate with 2 submarginal spines and
apical plumose setae; outer plate with apical
plumose setae. Maxilliped, left inner plate
with 2 strong apical teeth, right plate with
3; left outer plate with submarginal and api-
cal plumose setae; palp 4 articulate, dacty-
lus with two apical spines.
Gnathopod 1, basis elongate; posterior
lobe of carpus with 8 marginal setae; pro-
podus equal in length to carpus; palm with
submarginal setae, proximoposterior corner
with distinct spine. Gnathopod 2, basis
elongate; carpus with strong posterior con-
ical lobe with marginal setae; propodus ro-
bust and much larger than that of gnatho-
pod 1; palm with marginal spines and setae,
proximoposterior corner with tooth like
spine. Pereopods 3 and 4 similar in shape
and size; coxal plate of pereopod 4 su-
bquadrate and much larger than that of pe-
reopod 3; merus on both with distal anterior
lobe with 3 and 2 spines respectively. Pe-
reopod 5, approximately 75% the length of
pereopod 6; basis expanded posteriorly with
marginal and submarginal setae. Pereopod
7 slightly smaller than pereopod 6. Pereo-
pods 5-7 bases, anterior margins spinose,
posterior margins expanded posteriorly,
447
much stronger in pereopod 7; merus and
carpus of each distal posterior lobe spinose.
Pleopods 1—3 long and slender, pedun-
cles with 3 coupling hooks, except pleopod
3 with 2, rami with plumose setae. Uropod
1, peduncle with outer marginal with 4 bi-
furcate spines, both margins with distal
spine; inner ramus, with 2 marginal and 4
apical spines; outer ramus with 3 marginal
and 5 apical spines. Uropod 2, 75% the
length of uropod 1, peduncle outer margin
with 4 marginal spines, inner margin with
a strong distal spine; inner and outer rami
with 3 marginal and 4 apical spines. Uro-
pod 3, peduncle with 3 distal marginal and
1 submarginal spines; ramus with 7 apical
spines. Telson rounded, with 2 distal sub-
marginal spines and 5 small submarginal
setae.
Female (ovigerous).—All features same
as male except as noted below. Antenna 1,
27% of total body length; flagellum 7 artic-
ulate. Antenna 2, 63% of total body length
flagellum 14 articulate. Gnathopod 2 and
pereopods 3—5 with oostegites. Gnathopod
2, basis elongate; carpus longer than wide,
lacking posterior conical lobe and with 9
submarginal setae; palm and hind margin
distinct.
Etymology.—The specific epithet is a
name in apposition in reference to the first
author’s wife Sandra, for her support and
encouragement towards his research en-
deavors.
Material examined.—Male holotype,
4.80 mm, MCZ 25392, Travertine Spring,
approximately 1.9 km southeast of Texas
Spring, Death Valley National Park,
Inyo County, California, 36°26'28.40’N,
116°49'57.01"W, D. L. Threloff, 21 Dec
1997. Female (ovigerous) allotype, 3.36
mm, MCZ 25393, same data as holotype.
Male paratype, MCZ 25394, same data as
holotype, 4.88 mm. Female (ovigerous)
paratype, 3.36 mm, MCZ 25395, same data
as holotype. Male paratype, 4.80 mm, MCZ
25435, same data as holotype. Male para-
type, 4.72 mm, MCZ 25396, Texas Spring
outflow, 34 m downstream of discharge
448 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Ut
Fig. 3. Hyalella (Hyalella) muerta, male, 3.28 mm, USNM 230435.
point, Death Valley National Park, Inyo 25397, Texas Spring outflow, 13 m down-
County, California, 36°27'25.44"N, | stream of discharge point, Death Valley Na-
116°50'18.01"W, D. L. Threloff, 19 Dec tional Park, Inyo County, California,
1997. Male paratype, 5.20 mm, MCZ 36°27'25.86"N, 116°50'17.20"W, D. L.
VOLUME 113, NUMBER 2
449
Fig. 4. Hyalella (Hyalella) muerta, male, 3.28 mm, USNM 230435: Gnl, Gn2. Female, 3.24 mm, USNM
230436: Gnl, Gn2.
Threloff, 21 Dec 1997. Paratypes, USNM
230439, CASIZ 121605, same data as ho-
lotype. Paratypes, MCZ 25398, USNM
230411, CASIZ 121606, Texas Spring out-
flow, 34 m downstream of discharge point,
Death Valley National Park, Inyo County,
California, 36°27'25.44’N, 116°50'18.01’W,
D. L. Threloff, 19 Dec 1997. Paratypes,
MCZ 25399, USNM 230440, CASIZ
121607, Texas Spring outflow, 13 m down-
stream of discharge point, Death Valley Na-
tional Park, Inyo County, California,
IO LT 25.86 0 LIS 3017.20" WW,” D.*L.
Threloff, 21 Dec 1997.
Remarks.—Hyalella (H.) sandra is mor-
phologically similar to H. (H.) longicornis,
which is larger in size and is known from
only the type locality in Utah. Examination
450
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 5;
of the type specimens of H. (H.) longicornis
showed that the most significant differences
between H. (H.) sandra and H. (H.) longi-
cornis are: the ramus of uropod 3, H. (H.)
sandra robust, with 7 apical spines and H.
(H.) longicornis slender, with 4—5 apical se-
tae; the telson, H. (H.) sandra with 2 sub-
marginal spines and H. (H.) longicornis
with 2 long slender apical setae; the flagel-
lar articles of the male antenna 2, H. (H.)
sandra with 20—24 and H. (H.) longicornis
with 16; the palp of maxilla 1, H. (H.) san-
dra vestigial, 1 articulate and H. (H.) lon-
gicornis tall, length >2X the width; basis
of gnathopod 2 posterior margin, H. (H.)
sandra with 1—3 marginal setae and H. (H.)
longicornis with 9 marginal setae.
Discussion.—In 1874, Smith established
the genus Ayalella, describing Hyalella
dentata and HAyalella inermis, both from
Colorado. Faxon (1876), working on South
American hyalellids, regarded the genus AI-
lorchestes as the senior synonym of Hy-
Hyalella (Hyalella) muerta, male, 3.28 mm, USNM 230435.
alella and determined H. inermis was a va-
riety of H. dentata, calling it Allorchestes
dentata var. inermis. Nearly 30 years later,
Stebbing (1903) resurrected Hyalella and
established H. inermis as a valid species,
noting differences in the antennae, mouth-
parts, gnathopods and pereopods. Weckel
(1907), in re-examining the North Ameri-
can hyalellid species, concluded that H.
inermis, H. dentata and Hyalella faxoni
Stebbing, 1903 were all junior synonyms of
Hyalella knickerbockeri Bate, 1862 and
mentioned that only H. azteca var. inermis
lacked dorsal mucronations. Later, Barnard
(1958) provided a list of the Hyalella spe-
cies and listed H. faxoni and H. knicker-
bockeri as junior synonyms of H. azteca
and considered H. inermis a valid species.
However, Bousfield (1958, 1973) conclud-
ed that H. azteca is a single morphologi-
cally variable species with the number of
dorsal micronations varying from 1-3, that
specimens totally lacking dorsal mycrona-
VOLUME 113, NUMBER 2
Fig. 6. Hyalella (Hyalella) sandra, male, 4.88 mm
tions were H. azteca forma inermis, and
that both taxa occurred throughout the Unit-
ed States. Stevenson & Peden (1973) then
described HAyalella texana from Texas, a
species that coexisted with H. azteca.
Shortly thereafter, Cole & Watkins (1977)
described Hyalella montezuma from the
Montezuma Well system in Arizona, but
this species coexisted with H. azteca forma
inermis. Holsinger (1981) provided a list of
the 32 species of Hyalella and mentioned
that most workers agreed that H. dentata,
H. inermis and H. knickerbockeri are syn-
onyms of H. azteca. Lastly, Bousfield
(1996) described Ayalella (Ayalella) lon-
gicornis that lacked dorsal mucronations
and was known only from Utah [although
table 1 of Bousfield (1996) gives distribu-
tion as ““Texas’’].
Bousfield (1996) divided Hyalella into
three subgeneric groups (Hyalella, Austroh-
yalella, and Mesohyalella) based on geo-
graphical distributions and morphological
characters (i.e., body mucronations, the pro-
45]
, MCZ 25394.
podus of gnathopods 1 and 2, rami of uro-
pods 1 and 3, ornamentation of the telson).
The plesiomorphic subgenera Austrohyalel-
la and Mesohyalella are confined to conti-
nental South America while the more apo-
morphic subgenus Hyalella is endemic to
the West Indies, Central and North America
(Bousfield 1996). However, Bousfield
(1996) mentions that northern distributed
species of Mesohyalella show morphologi-
cal similarities to species in the North
American subgenus Hyalella.
Both species described here exhibit some
morphological characters similar to species
in the subgenus Mesohyalella [H. (H.)
muerta, smooth body, | plumose seta on the
inner plate of maxilla 2; H. (H.) sandra,
smooth body, 1 submarginal spine on the
inner plate of maxilla 2, spines on the tel-
son]. However, the presence of 5 sternal
gills, the morphology of male gnathopod 1
[H. (H.) muerta, propodus with 5 week and
short facial setae, palm margin convex and
short; H. (H.) sandra propodus with 5 week
452 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
AUS vy Y
ALAA AL
y iy! A} f
14%
By)
Wt,
v4 Mp
KY
Fig. 7. Hyalella (Hyalella) sandra, male, 4.88 mm, MCZ 25394.
and short facial setae, palm margin convex
and short, palmer angle with 1 short spine]
and the morphology of the female gnatho-
pod 2 of both species (propodus long and
slender) would place them in the subgenus
Hyalella. In addition, as both new species
lack copulatory spines on uropod 1, have a
peduncle and ramus of uropod 3 subequal
453
VOLUME 113, NUMBER 2
U1
Fig. 8. HAyalella (Hyalella) sandra, male, 4.88 mm, MCZ 25394: Al, A2, U3, T, Pll. Male, 4.80 mm, MCZ
ga4oa. U1, U2.
454 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 9. Hyalella (Hyalella) sandra, male, 4.88 mm, MCZ 25394: Gnl, Gn2. Female (ovigerous), 3.36 mm,
MEZ 25395: Gn, Gn2;
VOLUME 113, NUMBER 2
455
Fig. 10. Ayalella (Hyalella) sandra, male, 4.88 mm,
in length, and have setae on their telson
support their subgeneric placement. Lastly,
the North American distribution that defines
the subgenus Hyalella as suggested by
Bousfield (1996) are consistent with the
known distribution of the two new species
described here.
HAyalella (H.) muerta and H. (H.) sandra
occur in Death Valley National Park, Cali-
fornia but rarely if ever coexist together.
Hyalella (H.) muerta, the first North Amer-
ican hypogean hyalellid, is blind, lacks dor-
sal mucronations and antenna | is longer
than antenna 2. Hyalella (H.) sandra, col-
lected from epigean waters near Texas
Spring, also lacks dorsal mucronations but
has normal eye pigmentation and antenna |
is shorter than antenna 2, as in the other
North American Hyalella species. These
two new species bring the total number of
North American Hyalella taxa to seven.
Specimens of H. (H.) muerta were col-
lected in an artificial tunnel that was exca-
vated in the 1930’s. The tunnel was most
likely excavated in an effort to increase the
MCZ 25394.
volume of water that was being diverted as
a potable water supply. Prior to the devel-
opment of the tunnel, Texas Spring is be-
lieved to have issued water directly from
the local hillside. The interior of the tunnel
is typically 1.5 m wide by 1.5—2.0 m high
and is approximately 70 m in length. Flow
into the upstream portion of the tunnel de-
velops as water exits a fractured rock zone.
The water runs along the floor of the tunnel
in a stream that is 60—150 cm wide and 3—
30 cm deep. Overburden above the tunnel
consists of soft silt, 1-10 m thick. Collapse
of the tunnel has been prevented through
the installation of thick wooden cross mem-
bers that support the ceiling and side wall
surfaces of the tunnel. Specimens of H. (H.)
muerta were found among the submerged
roots originating from surface-inhabiting,
riparian plants. In 1995, a one meter portion
of the wooden tunnel structure collapsed. It
is not known if the concurrent sediment in-
put had any impact on the amphipod pop-
ulation.
At Texas Spring the water emerges from
456
local exposures of gravel and sand (Pistrang
& Kunkel 1958). The rate of flow has al-
ways been low, ranging from 0.2-0.5 cfs.
Because Texas Spring provides water for
human consumption, the water quality has
been regularly tested. Miller (1977) provid-
ed the following water chemistry analysis
for Texas Spring water: temperature: 31°C;
Silica: 25—40 mg/l; calcium 30 mg/l; sodi-
um: 150 mg/l; bicarbonate 330 mg/l; dis-
solved solids: 600—700 mg/l; pH: 7.5—8.5.
Specimens of H. (A.) sandra were col-
lected from Travertine Spring approximate-
ly 1.9 km southeast of Texas Spring and
13—34 m down stream from the Texas
Spring discharge. In mid to late 1970’s, the
potable water collection system at Texas
Spring was replaced, and the entire spring
flow was placed in a PVC pipe in an effort
to eliminate the percolation of water into
the ground. Between 1989 and 1994, Death
Valley National Park maintenance person-
nel diverted some of the piped water back
onto the ground in an effort to re-establish
a stream habitat. Maintenance personnel
then transplanted benthic sediment and veg-
etation from Travertine Spring to an area
down stream of Texas Spring tunnel with
the intentions of reinoculating the stream
with aquatic invertebrates and plants. Pre-
sumably, H. (H.) sandra was transported
with the sediments that established an in-
troduced population. Extensive sampling
has revealed one live specimen of H. (H.)
muerta occurring in the surface stream
downstream of Texas Spring tunnel. Prelim-
inary investigations therefore suggest that
H. (H.) muerta and H. (H.) sandra rarely
coexist in Death Valley National Park.
Although Death Valley is one of the dri-
est and hottest deserts in the New World,
the climate there has not always been so
harsh. During the Pleistocene the climate
was cooler and wetter, similar to that found
today around Lake Mono, 240 km (150 mi)
to the north. Numerous large pluvial lakes
occupied the many depressions in this area
and at that time Texas Spring would have
been at or only slightly above the shoreline
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
of pluvial Lake Manly. The aquatic com-
munities currently found in the springs and
streams of Death Valley are largely relicts
of these Pleistocene and earlier communi-
ties (Shepard 1992, 1993). Grayson (1993)
provides an excellent account of both Pleis-
tocene and Recent hydrology for the Death
Valley area.
In the desert southwest of the United
States, it appears that H. (H.) azteca has
been giving rise to new species via popu-
lations that have been isolated in thermally
constant waters. Thomas et al. (1994, 1997)
have proposed ecological isolation for the
species pair H. (H.) azteca and H. (H.)
montezuma, based on DNA and behavior.
Their behavioral studies led them to sepa-
rate two lineages of Hyalella in north-cen-
tral Arizona; swimmers that inhabit sub-
mersed vegetation in lakes and clingers that
inhabit springs dominated by emergent
macrophytes. Jackson (1912) also noted
two distinct locomotion behaviors. Hyalella
(H.) muerta and H. (H.) sandra both fall
into the clinger behavior category and like-
ly have speciated from local epigean pop-
ulations of H. (H.) azteca.
Another undescribed Hyalella (Hyalella)
species has also been found in Ash Mead-
ows National Wildlife Refuge, 45 km to the
east (in prep.). It also occurs in a warm
spring. Because of the discovery of a num-
ber of new species in such a small area, we
suggest that aquatic biologists more care-
fully collect and identify specimens in the
future, particularly when dealing with ther-
mally constant waters.
Acknowledgments
We thank Death Valley National Park for
permission to collect. We also thank M. Zu-
bowski (Royal Ontario Museum) for locat-
ing the types of H. (A). longicornis and C.
Serejo (Museu Nacional UF Rio De Janei-
ro) for the loan of H. caeca and help in
obtaining literature. The first author thanks
M. E Gable (Eastern Connecticut State Uni-
versity) for bringing this project to his at-
VOLUME 113, NUMBER 2
tention, E. A. Lazo-Wasem (Peabody Mu-
seum of Natural History, Yale University)
for helpful comments and discussion, and
A. B. Johnston (MCZ) for the use of the
departmental microscope.
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Barnard, J. L. 1958. Index to the families, genera and
species of the Gammaridean Amphipoda (Crus-
tacea).—Allan Hancock Foundation Publica-
tions, Occasional Paper Number 19:1—145.
Bate, C. S. 1862. Catalogue of the specimens of am-
phipodous Crustacea in the collection of the
British Museum, London, 399 pp.
Bousfield, E. L. 1958. Fresh-water amphipod crusta-
ceans of glaciated North America.—Canadian
Field Naturalist 72:55—113.
1973. Shallow-water Gammaridean amphi-
pods of New England. Cornell University Press,
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. 1996. A contribution to the reclassification of
neotropical freshwater hyalellid amphipods
(Crustacea: Gammaridea, Talitroidea).—Bollet-
tino del Museo Civico di Storia Naturale de Ve-
rona 20(1):175—224.
Cole, G. A., & R. L. Watkins. 1977. Hyalella monte-
zuma, a new species (Crustacea: Amphipoda)
from Montezuma Well, Arizona.—Hydrobiolo-
gia 52(2—3):175—-184.
Faxon, W. 1876. Exploration of Lake Titicaca by Al-
exander Agassiz and S. W. Garman. IV. Crus-
tacea.—Bulletin of the Museum of Comparative
Zoology 3:361—375.
Grayson, D. K. 1993. The desert’s past, a natural pre-
history of the Great Basin. Smithsonian Insti-
tution Press, Washington, 356 pp.
Holsinger, J. R. 1981. Amphipoda. Pp. 36—40 in S. H.
Hurlbert, G. Rodriguez & N. D. Santos, eds.,
Aquatic Biota of Tropical South America, Part
1: Arthropoda. San Diego State University, San
Diego, California.
Jackson, H. H. T. 1912. A contribution to the natural
history of the amphipod, Hyalella nickerbockeri
(Bate).—Bulletin of the Wisconsin Natural His-
tory Society 10(1—2):49-—60.
Miller, G. A. 1977. Appraisal of the water resources
of Death Valley California-Nevada. U.S. Geo-
457
logical Survey, Open File Report No. 77-728,
68 pp.
Pereira, V. EF G. C. 1989. Uma nova espécie de anfi-
pode cavernicola do Brasil-Hyalella caeca sp.
n. (Amphipoda, Hyalellidae)—Revista Brasi-
leira de Zoologia 6(1):49—55.
Pistrang, M. A., & E Kunkel. 1958. A brief geological
and hydrological reconnaissance of the Furnace
Creek wash area, Death Valley National Mon-
ument, California. United States Department of
Interior Groplogical Survey, Ground Water
Branch, 73 pp.
Ruffo, S. 1957. Una nuova specie troglobia di Hyalella
del Venezuela.—Annali del Museo Civico Di
Storia Naturale Genova 69:363—369.
Saussure, H. 1858. Mémoire sur divers crustacés nou-
veaux des Antilles et du Mexique.—Memoires
de la Societe Physique Histoire Naturelles Ge-
néve 14:417—496.
Shepard, W. D. 1992. Riffle beetles (Coleoptera: El-
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381.
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gered.—Aquatic Conservation: Marine and
Freshwater Ecosystems 3:351—359.
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ceans.—Annual Report of the United States
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Rica.—Proceedings of the United States Na-
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Stevenson, M. M., & A. E. Peden. 1973. Description
and ecology of Hyalella texana n. sp. (Crusta-
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Texas.—The American Midland Naturalist
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Thomas, P. E., W. Blinn, & P. Keim. 1994. A test of
an allopatric speciation model for congeneric
amphipods in an isolated ecosystem.—Journal
of the North American Benthological Society
13(1): 100-109.
1997. Genetic and behavioral divergence
among desert spring amphipod populations.—
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North America.—Proceedings of the United
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):458—464. 2000.
Caecidotea cumberlandensis, a new species of troglobitic isopod from
Virginia, with new records of other subterranean Caecidotea
(Crustacea: Isopoda: Asellidae)
Julian J. Lewis
217 W. Carter Avenue, Clarksville, Indiana 47129, U.S.A.
Abstract.—Caecidotea cumberlandensis is a subterranean asellid isopod de-
scribed from two caves in Cumberland Gap National Historic Park in south-
western Virginia. New records of several other subterranean Caecidotea are
also discussed: C. incurva, C. jordani, C. barri, C. paurotrigonus and C. ter-
esae. Comparison of the descriptions of C. paurotrigonus and C. dauphina
suggest that these species are conspecific.
In 1976 Dr. John Holsinger told me of a
subterranean isopod collected from Cliff
Cave in the Cumberland Gap area of south-
western Virginia that had been identified by
Fleming (1972a) as Asellus scrupulosus (a
troglophilic species with eyes and pigmen-
tation). Dr. Holsinger had related that this
identification might be in error since the
Cliff Cave specimen was eyeless and unpig-
mented, and suggested that I look at the
specimen if the opportunity presented. Flem-
ing’s identification was subsequently reject-
ed entirely by Holsinger & Culver (1988).
In 1997 I visited the Smithsonian Insti-
tution to assist in the sad task of curating
collections that remained in the office of my
mentor and friend, Dr. Thomas E. Bowman,
at the time of his death. During this process
I came upon the Cliff Cave specimen as
well as another vial of isopods from Indian
Cave, Lee Co., Virginia. Examination of
these specimens proved Dr. Holsinger’s sus-
picion correct, that the Cliff Cave speci-
mens represented a distinct taxon new to
science.
Family Asellidae G. O. Sars, 1897
Caecidotea Packard, 1871
Caecidotea cumberlandensis, new species
Figs. 1-3
Asellus scrupulosus.—Fleming, 1972a:
241.
Caecidotea species A.—Holsinger & Cul-
ver, 1988: 30-31, 37.
Material examined.—Virginia: Lee Co.,
Indian Cave, David A. Hubbard, Jr., 16 Mar
1993, 5.5 mm male holotype (USNM
291204), 4.5 mm male paratype, 6 female
paratypes (USNM 291205); Cliff Cave,
Russell M. Norton, 24 Nov 1966, 5.8 mm
male paratype (USNM 291206), including
a glass slide labelled “‘8-K A. scrupulosus”
and signed ““LEF”’ (L. E. Fleming) contain-
ing the first and second pleopods.
All specimens remain in the National
Museum of Natural History, Smithsonian
Institution.
Description.—Eyeless, unpigmented,
longest male 5.8 mm, female 4.7 mm; body
slender, about 5.2 as long as wide. Head
about 1.5 as wide as long, anterior margin
concave, postmandibular lobes moderately
produced. Pleotelson about 1.4 as long as
wide, sides subparallel, caudomedial lobe
moderately produced.
Mandibles with 4-cuspate incisors and la-
cinia mobilis, palp with rows of plumose
setae on distal segments. Maxilla 1 with 5
robust plumose setae on inner lobe, 13
spines on outer lobe. Antenna | reaching to
about mid-point of last segment of antenna
2 peduncle, flagellum of 6-7 segments, es-
thetes present on last 4 segments. Antenna
VOLUME 113, NUMBER 2
2, last segment of peduncle about 1.5
length of preceding segment, flagellum of
holotype with 40 segments.
Male pereopod 1, propus 2.6 as long as
wide, palmar margin with 2 large spines,
processes absent; female pereopod 1.2% as
long as wide, palmar margin similar to
male. Pereopods 2-7 similar, with moderate
setation, sexual dimorphism of pereopod 4
for grasping negligible, carpus about 2.2
as long as wide.
Male pleopod 1, protopod about 0.5
length of exopod, with 2-3 retinacula; ex-
opod about 1.8 as long as wide; lateral
margin slightly concave, distolateral margin
setae not plumose. Pleopod 2 exopod prox-
imal segment with 2 plumose setae, distal
segment with 3—5 plumose setae along mar-
gin. Endopod with rounded basal apophy-
Sis; tip with 2 distinct processes extending
subparallel to one another and approxi-
mately perpendicular to the axis of the en-
dopod: (1) lateral process subterminal, slen-
der, tapering slightly, and (2) cannula beak-
shaped. Pleopods 3—5, endopods present,
but unremarkable. Pleopod 3 exopod distal
margin with 3—4 short, non-plumose setae.
Pleopod 4 exopod, proximal setae absent;
sutures indistinct, suggestive of 2 barely
discernible, unconnected false sutures. Ple-
opod 5 exopod with faint transverse sutures.
Uropods of male about 0.5 X length of pleo-
telson, female similar.
Etymology.—The name refers to the
Cumberland Gap area in which the species
occurs. The suggested vernacular name is
the Cumberland Gap cave isopod.
Range.—This species is known only
from Cliff and Indian caves in the Cum-
berland Gap National Historic Park, Lee
Co., Virginia. Descriptions of the caves
were presented by Holsinger (1975). The
caves are about 500 meters apart, occur in
the same rock formation (Greenbrier Lime-
stone), and are probably disconnected parts
of the large Cudjos-Cumberland Saltpeter
Cave system (Holsinger, in litt.). The am-
phipod Stygobromus cumberlandus occurs
with C. cumberlandensis in Cliff Cave
459
(Holsinger 1978). The range of this amphi-
pod also includes Scott and Wise counties
in Virginia, suggesting the possibility of a
wider range for C. cumberlandensis.
Relationships.—Caecidotea cumberlan-
densis most closely resembles two other sub-
terranean species, C. bicrenata (northern Al-
abama to southern Illinois; Lewis 1982a)
and C. richardsonae (Tennessee and Virgin-
ia; Steeves 1963). These three species have
in common a male pleopod 2 endopod tip
with a terminal beak-shaped cannula and
subterminal lateral process, both extending
approximately perpendicular to the axis of
the endopod. These species are most easily
separated from one another by the shape of
the lateral process, which in C. cumberlan-
densis is thin and tapered distally, in C. bi-
crenata thicker and cylindrical, and in C. ri-
chardsonae finger-shaped and overlapped by
the cannula (the tip processes shown by
Lewis & Bowman 1977 for C. richardsonae
had been spread by pressure from a cover-
slip; the appearance depicted by Steeves
1963 is more typical). Differences and sim-
ilarities of key structures in these three spe-
cies are summarized in Table 1.
Caecidotea jordani (Eberly, 1966)
Material examined.—Indiana: Crawford
Co., seep spring flowing from bank of Blue
River at former site of Rothrock Mill, Wy-
andotte, Julian J. Lewis, Victor M. Lewis,
James J; Lewis, 30° Jul’ 1998, 5 males; 12
females; same locality, Julian J. Lewis, Sal-
isa T. Rafail, 1 Aug 1998, 4 males, 5 fe-
males.
Range.—The only previously known
population (a spring under Jordan Hall on
the campus of Indiana University, Bloo-
mington) was extirpated by termiticides.
The site reported above is a parafluvial hab-
itat where water seeps from a gravel bed
that extends under the floodplain of the
Blue River.
Caecidotea incurva (Steeves & Holsinger,
1968)
Material examined.—Tennessee: Blount
Co., Whiteoak Blowhole Cave, W. Reeves,
460 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1. Caecidotea cumberlandensis, 4.5 mm paratype male, Indian Cave, Lee Co., Virginia (a—b, d—h, j);
5.5 mm holotype male, same locality (c); 4.7 mm paratype female, same locality, (i): (a) habitus, (b) antenna
1, (c) antenna 2, (d) maxilla 1, outer lobe, (e) maxilla 1, inner lobe, (f) mandibular palp, (g) left mandible,
incisor and lacinia mobilis, (h) right mandible, incisor, (i) pleopod 2, (j) uropod.
VOLUME 113, NUMBER 2
461
Fig. 2.
mm male holotype, same locality, (b—c): (a) pereopod 1, (b) same, (c) pereopod 4, (d) same.
12 Aug 1999, 2 males, 2 females; Rich
Mountain Blowhole Cave, W. Reeves, 25
Aug 1999, 2 males, 1 female; Virginia:
Wythe Co., Early’s Cave, D. A. Hubbard,
Jr., 7 Jul 1997, 6 males, 4 females; Camp-
bell Cave, D. A. Hubbard, Jr., 14 Sep 1998,
1 male, 1 female; Lone Ash Cave #2, D. A.
Hubbard, Jr, 3 Nov 1997, 4 males, 13 fe-
males; Mockley Cave, D. A. Hubbard, Jr.,
29 Mar 1999, 1 male, 1 female.
Range.—This species was incompletely
described, but the endopod of the male sec-
ond pleopod is so distinct in appearance
(Steeves & Holsinger 1968) that a fairly
certain identification can be made. Holsin-
ger & Culver (1988) reported this species
in Virginia from McMullin Cave, Smyth
Co., and Groseclose Cave Number l,
Wythe Co. Other unpublished Virginia re-
cords identified and provided by J. R. Hol-
singer (in litt.) are Deep Spring and Bowles
Caecidotea cumberlandensis, 4.7 mm female paratype, Indian Cave, Lee Co., Virginia, (a, d); 5.5
Spring caves, Wythe Co., and Dolingers
Cave, Washington Co.
Caecidotea paurotrigonus (Fleming,
1972b)
Material examined.—Louisiana: St.
Mary Parish, holes dug in moist area at for-
est edge beside road, just outside entrance
to parish park at Burn’s Point, at end of
state highway 317, coll. D.W.D., 23 Aug
1981, 1 male, 1 fragment, 9 females.
Range.—This groundwater species was
previously known from a single male de-
scribed by Fleming (1972b) from a ditch in
southwestern Mississippi. The new locality
is the first report of a subterranean asellid
from Louisiana. Identification of C. pau-
rotrigonus is obscured by Fleming’s de-
scription. Based on a single 16.7 mm male,
the description provided drawings of key
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3:
Caecidotea cumberlandensis, 4.5 mm paratype male, Indian Cave, Lee Co., Virginia (a—c, e—g); 5.8
mm male, Cliff Cave, Lee Co., Virginia (d): (a) pleopod 1, (b) pleopod 2, (c) same, endopod tip, (d) same, (e)
pleopod 3 exopod, (f) pleopod 4 exopod, (g) pleopod 5 exopod.
structures, but the quality of the drawings
and the interpretation of the structures were
both questionable. Modlin (1986) on the
other hand provided a detailed description
of Caecidotea dauphina based on a 7.5 mm
male. Unfortunately, three of the four struc-
tures figured by Fleming for C. paurotri-
gonus bear a strong resemblance to that of
C. dauphina: (1) pereopod 1 propods of
very similar dimensions, margin in C. dau-
phina with basal spine, medial and distal
processes, identical in C. paurotrigonus ex-
VOLUME 113, NUMBER 2
463
Table 1.—Comparison of selected structures of male C. cumberlandensis, C. bicrenata and C. richardsonae
useful for separating the species.
C. cumberlandensis
C. bicrenata C. richardsonae
Pereopod 1 propod
palmar margin processes absent
Pleopod |
distolateral lobe absent
Pleopod 2 exopod 3-5 setae
Pleopod 2 endopod
lateral process
Proportion of uropod
length to pleotelson 0.5 length
cept the spine is replaced by a process (as
typical of more mature specimens); (2) ple-
opod 1 with essentially identical shapes, se-
tation patterns and quantity of retinacula;
(2) pleopod 2 endopod tips with short api-
cal cannula directed mediad, and short, un-
remarkable knob-like lateral process. I sus-
pect that C. dauphina represents a well de-
scribed juvenile of the poorly described, but
conspecific C. paurotrigonus. Unravelling
this will require redescription of C. pauro-
trigonus and perhaps more male specimens
for comparison from Mississippi and Ala-
bama.
Caecidotea barri (Steeves, 1965)
Material examined.—Kentucky: Wood-
ford Co., small spring 0.6 mile E. Clifton,
in side valley above waterfall, elevation
about 670 feet, Julian J. Lewis, Victor M.
Lewis, James J. Lewis, 19 Feb 1995, 3
males, 4 females.
Range.—This species was previously
known only from the type-locality, Clifton
Cave, Woodford Co., Kentucky (Steeves
1965), which was bulldozed shut by high-
way workers. The new locality is across the
valley from Clifton Cave.
Caecidotea teresae Lewis, 1982b
Material examined.—Indiana: Floyd Co.,
well on Grant Line Road, Julian J. Lewis,
26 Apr 1995, 1 male, 1 female.
thin, tapered distally
present absent
absent present
12-15 setae 12—15 setae
cylindrical, not tapered long digitiform
1.5—2.0 length 1.5—2.0 length
Range.This species was apparently extir-
pated by termiticides at the two previously
known localities on the campus of Indiana
University Southeast, New Albany. The
well from which the new specimens were
taken is found in a pasture across Grant
Line Road from the I.U.S. campus. Accord-
ing to the owner, the well was hand dug at
the end of the 19th century and was 21 feet
deep, with 8 feet in soil and the bottom 13
feet through New Albany Shale.
Acknowledgments
The description of Caecidotea cumber-
landensis was prepared during a visit to the
National Museum of Natural History fund-
ed by a Smithsonian visiting scientist grant.
Caecidotea jordani was rediscovered dur-
ing the bioinventory of Blue River area
caves funded by a Rodney Johnson/Kathar-
ine Ordway Stewardship Endowment Grant
from The Nature Conservancy. Caecidotea
teresae was rediscovered during field work
funded by the Indiana Non-game and En-
dangered Wildlife Program. Caecidotea
barri was found during field work funded
by the U.S. Fish & Wildlife Service and
Kentucky Nature Preserves Commission
(Dr. Thomas C. Barr, Jr. principle investi-
gator). I thank Dr. John R. Holsinger (Old
Dominion University), Dr. Brian Kensley
(Smithsonian Institution), Dr. Guy Magniez
(University of Dijon), Mr. David Hubbard,
464
Jr. (Virginia Division of Mineral Resourc-
es), and Mr. Allen Pursell (The Nature Con-
servancy) for reading the manuscript and
making suggestions on its improvement.
Literature Cited
Eberly, W. R. 1966. A new troglobitic isopod (Asel-
lidae) from southern Indiana.—Proceedings of
the Indiana Academy of Science 75:286—288.
Fleming, L. E. 1972a. The evolution of the eastern
North American isopods of the genus Asellus
(Crustacea: Asellidae).—International Journal
of Speleology 4:221—256.
. 1972b. Four new species of troglobitic asel-
lids (Crustacea: Isopoda) from the United
States.—Proceedings of the Biological Society
of Washington 84:489—499.
Holsinger, J. R. 1975. Descriptions of Virginia
Caves.—Virginia Division of Mineral Resourc-
es, Bulletin 85, 450 pp.
. 1978. Systematics of the subterranean amphi-
pod genus Stygobromus (Crangonyctidae), Part
II: Species of the eastern United States.—
Smithsonian Contributions to Zoology 266:1—
144.
, & D. C. Culver. 1988. The invertebrate cave
fauna of Virginia and a part of eastern Tennes-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
see: Zoogeography and ecology.—Brimleyana
14:1-162.
Lewis, J. J. 1982a. Systematics of the troglobitic Cae-
cidotea (Crustacea: Isopoda: Asellidae) of the
southern Interior Low Plateaus.—Brimleyana 8:
65-74.
. 1982b. A diagnosis of the Hobbsi Group, with
descriptions of Caecidotea teresae, new spe-
cies, and C. macropropoda Chase and Blair
(Crustacea: Isopoda: Asellidae).—Proceedings
of the Biological Society of Washington 95:
338-346.
, & T. E. Bowman. 1977. Caecidotea caroli-
nensis, n. sp., the first subterranean water slater
from North Carolina (Crustacea: Isopoda: Asel-
lidae).—Proceedings of the Biological Society
of Washington 90:968—974.
Modlin, R. E 1986. Caecidotea dauphina, a new sub-
terranean isopod from a barrier island in the
northern Gulf of Mexico (Crustacea: Isopoda:
Asellidae)—Proceedings of the Biological So-
ciety of Washington 99:316—322.
Steeves, H. R., III. 1963. The troglobitic asellids of the
United States: The Stygius Group.—American
Midland Naturalist 69:470—48 1.
. 1965. Two new species of troglobitic asellids
from the United States——American Midland
Naturalist 73:81—84.
, & J. R. Holsinger. 1968. Biology of three new
species of troglobitic asellids from Tennes-
see.—American Midland Naturalist 80:75-—83.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):465—479. 2000.
Euphilomedes cooki, a new species of myodocopid ostracode from
Moreton Bay, SE Queensland, Australia
Elizabeth Harrison-Nelson and Louis S. Kornicker
Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A.
Abstract.—A new species of myodocopid ostracode in the subfamily Phi-
lomedinae from Moreton Bay, S.E. Queensland, Australia, is described and
illustrated. A key is presented to the ten known species of Euphilomedes having
primary furcal claws 1, 2, 4, and 6.
This work describes and illustrates a new
species of myodocopid ostracode from
Moreton Bay, S.E. Queensland, Australia,
which is north of Brisbane, near the south-
ern end of the Great Barrier Reef.
Disposition of specimens.—The holotype
and paratypes have been deposited at the
Queensland Museum (QM), South Bris-
bane, Queensland.
Superorder Myodocopa Sars, 1866
Order Myodocopida Sars, 1866
Suborder Myodocopina Sars, 1866
Family Philomedidae Miiller, 1906
This family contains two subfamilies,
Philomedinae Miiller, 1906, and Pseudo-
philomedinae Kornicker, 1967, both with
representatives in the vicinity of S.E.
Queensland, Australia (Kornicker 1994).
Philomedinae Miiller, 1906
This subfamily includes eight genera:
Philomedes Liljeborg, 1853, Pleoschisma
Brady, 1890, Scleroconcha Skogsberg,
1920, Paraphilomedes Poulsen, 1962, Eu-
philomedes Kornicker, 1967, Anarthron
Kornicker, 1975, Igene Kornicker, 1975,
and Zeugophilomedes Kornicker, 1983. A
new species of Euphilomedes is described
herein.
Euphilomedes Kornicker, 1967
Type species.—Euphilomedes nodosa
Poulsen, 1962 (subsequent designation by
Kornicker 1967).
Composition.—Including the new spe-
cies described herein this genus includes 19
species plus one subspecies (Kornicker
19912371 995e16).
Distribution.—The genus is cosmopoli-
tan except in Arctic and Antarctic waters.
The known depth range is shallow water to
2250 m, but members have been collected
mostly from the continental shelf and upper
slope (Kornicker & Harrison-Nelson 1997:
14). Three species of the genus have been
reported previously from the vicinity of
Australia: E. corrugata (Brady 1897) from
off Port Jackson and in Flinders Passage at
depths of 3.7—18.3 m, E. walfordi Poulsen,
1962, from the Coral Sea at a depth of 50
m, and E. erynx Kornicker, 1995, from the
continental slope off New South Wales at a
depth of 220 m. The new species described
herein is from Middle Banks, northern
Moreton Bay, S.E. Queensland, Australia,
at a depth of 15—25 m.
Euphilomedes cooki, new species
Figs. 1-9
Etymology.—The species is named in
honor of Stephen Cook formerly of the
Queensland Museum, Australia, who col-
lected some of the specimens.
466
Holotype.—Queensland Museum W2492,
undissected adult female in alcohol.
Type locality.—Miiddle Banks, northern
Moreton Bay (27.02°S, 153.25°E), Queens-
land, Australia, about 20 km offshore from
the mainland, depth 15-25 m. All speci-
mens part of Queensland Museum Registra-
tion Number 11879.
Paratypes.—12 adult females; 3 A-1
males; 1 A-1 female.
Distribution.—Middle Banks, northern
Moreton Bay, S.E. Queensland, depth 15-—
25 m, clean sand or sand and shell. Col-
lected with Smith-MclIntyre grab in Novem-
ber 1983 and November 1984.
Description of adult female (Figs. 1-7a—
7c).—Carapace elongate with shallow in-
cisur (Fig. la). Left valve extends past right
valve along free margin. Posterodorsal
hinge area straight and without valve over-
lap.
Ornamentation: Carapace with pits of
various sizes and shapes. Each pit with nar-
row raised border. Some pits two or three
times larger than others; pits larger in pos-
terior half of valve. Valve edge with long
single bristles. Long single bristles sparsely
distributed on valve surface.
Infold: Rostral and anteroventral infold
(Fig. 1b,c,e, 7a), infold of caudal process
and posteroventral infold (Fig. 1d,f, 7b),
each with row of bristles. Anteroventral in-
fold with several parallel ridges (Fig. 1c).
Selvage: Broad lamellar prolongation of
right valve in vicinity of rostrum with long
marginal hairs; lamellar prolongation nar-
rower and with shorter marginal hairs along
ventral margin. Lamellar prolongation of
left valve obscured.
Central adductor muscle attachments
(Fig. 1g): About 30 attachments on each
side. These represented by large pits on
each valve just anterior to midlength; most
pits ventral to midheight.
Carapace Size (mm): Average length
1.89, range 1.81—1.95; average height 1.36,
range 1.29-1.42; average height 72% of
length, range 71-74; n = 11.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Height
Length Height % length
Holotype 1.94 39 TW
Paratypes 1.84 i Wee) 74
25 1.40 93
LoS 1.42 73
87 Sy, 71
1.81 Vol 71
Peo 136 A
oS 1.41 2
1.86 i335 73
87 1has) 7 71
1.81 29 7
First antenna (Fig. 2a): 1st joint with dis-
tal medial spines forming rows. 2nd joint
with dorsal, ventral, and lateral spines, and
3 bristles (1 dorsal, 1 ventral, 1 medial). 3rd
joint short with 3 bristles (2 dorsal, 1 ven-
tral). 4th joint with 6 bristles (2 dorsal, 4
ventral). Sth joint about same length as 4th
joint, with long sensory bristle with about
7 short marginal filaments. Long medial
bristle of 6th joint with base near dorsal
margin and with long proximal and short
distal spines. 7th joint: a-bristle similar to
bristle of 6th joint; b-bristle with 2 fila-
ments near midlength, 2 subterminal fila-
ments, and bifurcate tip; c-bristle long with
marginal filaments. 8th joint: d- and e-bris-
tles long, bare, and with blunt tips; f- and
g-bristles long with marginal filaments. Not
all filaments of bristles of 7th and 8th joints
shown in illustration.
Second antenna: Protopodite bare. En-
dopodite 2- or 3-jointed (Fig. 2b): 1st joint
with row of 5 short bare proximal bristles
and 1 very long spinous distal bristle; 2nd
joint with very long spinous proximal bris-
tle (bristle 2 % times length of long bristle
of Ist joint) and short bare terminal bristle
(the terminal bristle could be interpreted to
be on a 3rd joint). Exopodite: Ist joint with
minute straight medial terminal bristle; bris-
tle of 2nd joint reaching well past 9th joint,
with few dorsal spines and abundant short,
fairly stout, ventral spines, no natatory
hairs; bristles of 3rd and 4th joints with
short slender distal spines, no natatory
VOLUME 113, NUMBER 2 467
a
Fig. 1. Euphilomedes cooki specimen 12, adult female paratype: a, outline of complete specimen from left
side, length 1.84 mm, note four epibionts attached to edge of valve. b—d, interior views of left valve: b, anterior
end; c, anteroventral margin; d, posterior end. e, f, interior views of right valve: e, anteroventral margin; f,
posteroventral margin. g, ends of central adductor muscles projecting from right side of body, anterior toward
right. Abbreviation are: ant: antenna; Bo: Bellonci organ; cx: coxale; e: edge of valve; end: endopodite; ex:
exopodite; epip: epipodite; fu: furca; gen: genitalia; gird: girdle; im: inner margin of infold; li:list; lv: lateral
view; me: medial eye; mnd: mandible; mv: medial view; mx: maxilla; nabs: not all bristles shown; prot: pro-
topodite; ul: upper lip; Y-scl: Y-sclerite. Roman numerals designate endites. Arrow on illustration indicates
anterior.
468
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
APACE
ee = NAN ON
oN
7
Vi
\
/
\ prot
\
7
- Y
Lh \
Z.
\ Vi
\ \ y
SY ‘
Riga:
Euphilomedes cooki specimen 12, adult female paratype: a, right Ist antenna, lv; b, left 2nd antenna,
endopodite and distal part of protopodite, mv; c, right 2nd antenna, exopodial joints 8 and 9, only proximal
parts of some bristles shown, lv; d, right furcal lamella, lv; e, left lateral eye, lv; f, posterior of body from right
side; g, outline of unextruded egg, drawn at same scale as “‘f’’. (For abbreviations see Fig. 1.)
VOLUME 113, NUMBER 2
hairs; bristles of joints 5—8 longer than bris-
tles of joints 2—4, with natatory hairs, no
spines; 9th joint with 7 bristles (4 long and
2 short with natatory hairs, 1 very short and
with short marginal spines (Fig. 2c, only
proximal parts of some bristles shown);
joints 2—8 with row of terminal spines.
Mandible (Fig. 3a, b): Coxale endite spi-
nous, tip bifurcate, with small ringed bristle
near base. Basale: medial surface and ven-
tral and dorsal margins with rows of spines;
medial surface with 5 bristles in proximal
ventral corner (3 pectinate unringed, 2
ringed and with long proximal and short
distal spines), and 1 short ringed bristle
closer to midlength with long proximal and
short distal spines; dorsal margin with 3
bristles (1 near midlength, 2 terminal); 7
bristles with long proximal and short distal
spines present on or near ventral margin (2
longest bristles distal and with bases on
ventral margin; 5 shorter bristles with bases
slightly lateral). Exopodite slightly more
than % length of dorsal margin of Ist en-
dopodial joint, with distinct distal hirsute
pad and few terminal spines, and 2 bristles
(outer bristle with short marginal spines and
about % length of inner bristle; inner bristle
reaching midlength of 2nd endopodial joint,
with long spines near midlength and short
distal spines). 1st endopodial joint with me-
dial spines and 4 ventral bristles. 2nd en-
dopodial joint: dorsal margin with 2 long
bristles in proximal group and 5 spinous
bristles in distal group (2 long with basis
on margin; 3 short with bases medial (prox-
imal 2 with long spines, distal 1 with short
spines)); ventral margin with bristles in 2
groups (2 in proximal group, 3 in distal
group); medial surface with spines forming
rows. 3rd endopodial joint with 3 pectinate
claws (dorsal claw short), and 3 ringed bris-
tles.
Maxilla (Figs. 3b, 4a—c): Precoxale and
coxale with fringe of long hairs. Coxale
with plumose dorsal bristle. Basale with 3
distal bristles. Exopodite with 3 bristles
(proximal short bristle bare, long middle
bristle with long spines, other long bristle
469
with short spines). Ist endopodial joint with
1 alpha-bristle and 4 beta-bristles. 2nd en-
dopodial joint with 3 pectinate claw-like
bristles and about 7 ringed bristles. Endites
with stout spinous and pectinate bristles
(endite I with 9 bristles, endite II with 6
bristles, endite III with 8 bristles).
Fifth limb (Figs. 4d—f, 5): Epipodite with
46 bristles. Endite I with 5 bristles (only 3
shown in illustration); endite II with 6 bris-
tles; endite III with 8 bristles. Ist exopodial
joint: anterior side with 2 bristles (with long
spines) on distal edge; outer corner with 2
small slender bristles with few marginal
hairs; main tooth with proximal peg fol-
lowed by 3 pointed teeth (teeth worn down
on specimen 12) and 1 large squarish tooth;
ringed spinous bristle proximal to teeth. 2nd
exopodial joint: posterior side with 1 prox-
imal bristle and group of 3 distal bristles
(middle bristle long, others short (short
bristle obscured on right limb of specimen
12)). 3rd exopodial joint: inner lobe with 3
bristles, outer lobe with 2 bristles with long
spines. Fused 4th and 5th exopodial joints
with 8 bristles.
Sixth limb (Figs. 5a, 6a): Epipodite with
3 spinous bristles. Endite I with 3 bristles;
endite II with 4 bristles; endites III and IV
each with 9 bristles. End joint with spines
and hairs and 18 bristles (most with long
proximal hairs and short distal spines).
Seventh limb (Fig. 6b, c): Each limb with
13-15 bristles: proximal group with 8—10
bristles (specimen 12: 5 or 6 on peg side,
4 on comb side; specimen 4 (right limb), 5
on peg side, 3 on comb side), each bristle
with 3—5 bells and marginal spines; termi-
nal group with 5 bristles (3 on peg side, 2
on comb side). Each bristle with 3—7 bells
and marginal spines. Terminal comb with
12 alate teeth; 2 small curved pegs (with
proximal teeth) present opposite comb.
Furca (Fig. 2d, f): Each lamella with 11
claws: claws 1, 2, 4, and 6 primary; claws
3, 5, 7-11 secondary. Primary claws with
stout posterior teeth; teeth of claw 1 stouter
than teeth of other primary claws and with
stout medial teeth; secondary claws with
470 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
a
sw eSee
I a
—— OTE oTo—E—E=x=«: oo —
Fig. 3. Euphilomedes cooki specimen 12, adult female paratype: a, right mandible, mv; b, left maxilla and
mandible in place on body, nabs, mv; c, medial eye and Bellonci organ from left side. (For abbreviations see
Fig. 1.)
VOLUME 113, NUMBER 2 471
Fig. 4. Euphilomedes cooki specimen 12, adult female paratype: a—c, right maxilla: a, complete limb, lv; b,
bristle of distal end of Ist exopodial joint, mv; c, bristles and claws of 2nd endopodial joint, mv. d—f, left 5th
limb, endites I, II, and III, respectively. (For abbreviations see Fig. 1.)
472 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
An
Fig. 5. Euphilomedes cooki specimen 12, adult female paratype: a, upper lip, left 6th limb, and right 5th
limb in place on body, not all bristles shown. b, c, posterior views of distal end of right fifth limb, respectively.
d, e, posterior and anterior views of distal end of left fifth limb, respectively. (For abbreviations see Fig. 1.)
marginal spines. Right lamella with few Bellonci organ (Fig. 3c): Elongate with
distal anterior spines and also spines medial short wide part (bearing striations) near
and proximal to claw 1. Right lamella an- midlength and narrow tapered tip bearing
terior to left by % width of claw 1. minute spines.
VOLUME 113, NUMBER 2 473
\
\
)
kk
Ly
YE
Gp
\
}
\
~
CUP IT/
p
i
Fig. 6. Euphilomedes cooki specimen 12, adult female paratype: a, right 6th limb, lv; b, right 7th limb; c,
tip of left 7th limb, only proximal part of bristles shown. (For abbreviations see Fig. 1.)
474
Eyes: Medial eye with brown pigment
(Fig. 3c). Lateral eye smaller than medial
eye, unpigmented, with 4 divided amber-
colored ommatidia (Fig. 2e).
Upper lip (Figs. 5a, 7c):-Projecting
slightly anteriorly, with anterior and lateral
glandular processes.
Genitalia (Fig. 2f): Small oval.
Anterior of body: Convex.
Posterior of body (Fig. 2f): Evenly
rounded, bare.
Y-sclerite (Fig. 2f): With ventral branch.
Number and lengths of eggs: Specimen
12 with about 16 unextruded eggs (Fig. 2g).
Specimen 4 with 18 eggs in marsupium
(lengths of 2 eggs 0.406 mm, 0.408 mm).
Gut content: Specimen 12 with unrec-
ognizable amber-colored particles in gut.
Epizoa: Specimens 12 and 4 with vase-
shaped protistans along posterior margin
(Fig. la, f).
Description of A-1 male (Fig. 7d—f).—
Carapace similar in shape and ornamenta-
tion to that of adult female.
Carapace size (length, height in mm):
1.59, 1.06, height 67% of length (specimen
13); 1.58, 1.13, height 72% of length.
Second antenna: Protopodite bare with
narrow slightly curved proximal pivotal
sclerite. Endopodite 3-jointed (Fig. 7d): Ist
joint short with 4 small proximal ringed
bristles and 1 distal long stout ringed bristle
(broken off in illustrated endopodite (Fig.
7d)); 2nd joint elongate with 3 small ringed
bristles; 3rd joint elongate with 2 small
ringed terminal bristles with short spines.
Exopodite 9-jointed: Ist joint with small
medial terminal spine; bristle of 2nd joint
less than twice length of exopodite, with
spines along ventral edge; bristles of joints
3-8 fairly short, longer than bristle of 2nd
joint, but less than twice length of exopod-
ite, with ventral spines stouter than those of
bristle of 2nd joint, without natatory hairs;
9th joint with 6 bristles (2 short lateral at
distal dorsal corner, 4 longer terminal (lon-
gest at ventral end and less than twice
length of exopodite)), all with marginal
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
spines, without natatory hairs; joints 2-8
with row of spines along terminal edges.
Fifth limb (Fig. 7e, f): Similar to that of
adult female.
Sixth limb: With 3 epipodial bristles. En-
dite I with 3 bristles; endite II with 4 bris-
tles; endite III with 8 bristles.
Seventh limb: Limb with 12 tapered bris-
tles with marginal spines: proximal group
with 7 bristles (3 on comb side, 4 on peg
side), each with 2 or 3 bells; terminal group
with 5 bristle (2 on comb side, 3 on peg
side), each with 2—5 bells. Terminus with
comb of about 11 alate teeth opposite 2
small pegs.
Furca: Similar to that of adult female ex-
cept with only 4 secondary claws following
primary claw 6.
Lateral eye: Well developed with many
ommatidia and black pigment between
them.
Description of A-1 female (Figs. 8,9).—
Carapace similar in shape and ornamenta-
tion to that of adult female, but narrower.
Carapace size (length, height in mm):
1.58, 1.02; height 65% of length (specimen
1):
Remarks.—The absence of natatory hairs
on bristles of the exopodite of the 2nd an-
tennae of A-1 males and females indicates
that the juveniles of this species are inca-
pable of efficient swimming.
Comparisons.—Each lamella of the furca
of the new species E. cooki bears primary
claws 1, 2, 4, and 6, secondary claws 3 and
5, and 5 additional secondary claws follow-
ing primary claw 6. Only one species of
Euphilomedes having a similar distribution
of primary and secondary claws has been
described previously from the vicinity of
Australia, E. walfordi, which is known from
only the adult male, and so is not directly
comparable in all morphological characters
to the adult female and A-1 instar male of
E. cooki described herein. However, the fol-
lowing differences between the adult male
of the E. walfordi and the adult female of
E. cooki are considered significant: 1, The
carapace length of the adult female cooki is
VOLUME 113, NUMBER 2 475
Fig. 7. Euphilomedes cooki specimen 2, adult female paratype: a, b, interior views of anterior and posterior
ends of right valve, respectively; c, upper lip, anterior toward left. d, specimen 13, A-1 male paratype, left 2nd
antenna, endopodite and distal part of protopodite, mv. e, f, right 5th limb, av: e, distal end endite I. (For
abbreviations see Fig. 1.)
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
400M 20KY 200HM 20K¥
200HM OO = 190M ZOKY
Fig. 8. Euphilomedes cooki, A-1 female paratype (specimen 1), left valve: a, lateral view of valve, length
1.58 mm; b, dorsal view of valve, anterior to left; c, anterior view of valve, ventral to right; d, anteroventral
margin of valve, lv, from ‘‘a’’; e, area in vicinity of central adductor muscle, from ‘‘a’’; f, detail of reticulations,
from lower left part of ‘‘e’’.
VOLUME 113, NUMBER 2 477
——20FM 20K¥~—s«o
7 *
PE
ek
00
40PM 2OKY¥
10rM
Fig. 9. Euphilomedes cooki A-1 female paratype (specimen 1), left valve: a, detail of pits and bristles in
upper right part of Fig. 8a; b, detail of bristle shown in upper right of ‘‘a’’; c, detail of bristle and reticulations
in posterior part at midheight of Fig. 8a; d, detail of bristle and reticulations in anterior part at midheight of
Fig. 8a; e, detail of bristles and reticulations near incisure in Fig. 8d.
478
1.73—1.95 mm compared to 3 mm for the
adult male walfordi; 2, the outer surface of
the carapace of the adult female cooki has
few hairs, whereas that of the adult male
walfordi, according to Poulsen (1962:371),
‘‘is densely covered with short hairs’’. The
seventh limb of the adult male walfordi has
19 bristles without marginal spines (Poul-
sen 1962:372), whereas that limb of the
adult female cooki has 13-15 bristles with
marginal spines (when differences occur in
the number of bristles on the 7th limbs of
male and female myodocopids, the male
generally has fewer bristles than the fe-
male).
The key to species of Euphilomedes pre-
sented below is restricted to those species
having furcal lamellae with primary claws
1, 2, 4, and 6, and secondary claws 3, 5,
and 7+.
Key to species of Euphilomedes having
furcal claws 1, 2, 4, and 6 primary*
1. Lower angle of posterior shell margin
withosmialletoothr nce ca%\ tears sinc ie oe
E. sinister (Kornicker, 1974)
1. Lower angle of posterior shell margin
a Xe: <0) 1e) .0), Xe) ware.
VALCDOUE COOUME ona ex chins Ste Gk ae 2
2. Seventh limb with less than 12 cleaning
DEISCIES Se ean ee ner oe one ae cere 3
2. Seventh limb with 13-15 cleaning bris-
tles E, cooki, 1. sp.
2. Seventh limb with more than 15 cleaning
bristles E. walfordi (Poulsen, 1962)
3. 2nd joint of endopodite of female 2nd
antenna with only the long plumose bris-
(1G 3 secret one ee E. africana (Klie, 1940)
3. This joint with an additional distal bristle
Late CSS OUND OMe iO! On Lh Oe OE et
oe © ee
4. The distal bristle is only a spine, shorter
than: the width of- the jomt 5733, @. 2.
E. japonica (Miller, 1890)
4. This bristle slightly longer than width of
(0.1 11 Cees emer eer 17 SSR Atop tum racer ae See oe)
E. morini (Kornicker & Harrison-Nelson,
1997)
4. This bristle longer than twice the width
ol the 4oinbos: ec eae es ote 5
5. This bristle is placed dorsally on the
joint E. nodosa (Poulsen, 1962)
© #0) @ 8) @) 0 a 10
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
5. This bristle is placed on the narrow tip
Of thé joint. o> 62 eee 6
6. Furca with more than 12 pairs of claws
E. smithi (Poulsen, 1962)
6. Furca with 11 or less pairs of claws ... 7
7. Shell with a dorso-posterior, chitinized
PROCESS: ar E. producta (Poulsen, 1962)
7. Shell without such process
E. longiseta (Juday, 1907)
©) 6% 'e, (0° <6) ‘0; 6) (0: 0) she) <0 Ke,
* Key adapted in part from the key to spe-
cies of the genus Euphilomedes in Poul-
sen (1962:362).
Acknowledgments
Collections of specimens from Moreton
Bay described herein were made by P.
Saenger and S. Cook, Queensland Museum,
Australia. We thank Matthew Kane, Smith-
sonian Institution (SI) volunteer, for inking
penciled camera lucida drawings of append-
ages, Walter Brown (SI) for SEM micro-
graphs of the carapace, and Andrew R.
Parker, University of Oxford, for reviewing
the manuscript.
Literature Cited
Brady, G. S. 1890. On Ostracoda collected by H. B.
Brady, Esq., L.L.D.ER.S., in the South Sea Is-
lands.—Transactions of the Royal Society of
Edinburgh, 35(part 2):489—525.
. 1897. A supplementary report on the crusta-
ceans of the group Myodocopa obtained during
the Challenger Expedition, with notes on other
new or imperfectly known species.—Transac-
tions of the Zoological Society of London 14:
85-100.
Juday, C. 1907. Ostracoda of the San Diego region, II.
Littoral forms.—University of California Pub-
lications in Zoology 3:135—156.
Klie, W. 1940. Beitrage zur Fauna Eulitorials von
Deutsch Stidwest-Afrika, II: Ostracoden von der
Kiiste Deutsch-Stidwest-Afrika.—Kieler Meer-
esforschungen 3:403—448.
Kornicker, L. S. 1967. The myodocopid Ostracod fam-
ilies Philomedidae and Pseudophilomedidae
(New Family).—Proceedings of the United
States National Museum 120(3580):1-—35.
. 1974. Revision of the Cypridinacea of the
Gulf of Naples (Ostracoda).—Smithsonian
Contributions to Zoology 178:1—64.
1975. Antarctic Ostracoda (Myodocopina)
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Parts 1 and 2.—Smithsonian Contributions to
Zoology 163:1—720.
. 1983. Zeugophilomedes, a new genus of my-
odocopine Ostracode.—Proceedings of the Bi-
ological Society of Washington 96:478—480.
. 1991. Myodocopid Ostracoda of hydrothermal
vents in the Eastern Pacific Ocean.—Smithson-
ian Contributions to Zoology 516:1—46.
. 1994. Ostracoda (Myodocopina) of the SE
Australian continental slope, Part 1.—Smith-
sonian Contributions to Zoology 553:1—200.
. 1995. Ostracoda (Myodocopina) of the SE
Australian Continental Slope. Part 2.—Smith-
sonian Contributions to Zoology 562:1—97.
, & E. Harrison-Nelson. 1997. Myodocopid Os-
tracoda of Pillar Point Harbor, Half Moon Bay,
California.—Smithsonian Contributions to Zo-
ology 593:1-53.
Liljeborg, W. 1853. Ostracoda. Pp. 92—130 in De Crus-
479
taceis ex ordinibus tribus: Cladocera, Ostracoda
et Copepoda in Scania Occurrentibus. Lund:
Berlingska Boktryckeriet.
Miiller, G. W. 1890. Neue Cypridiniden.—Zoologische
Jahrbuecher 5:211-—252.
1906. Die Ostracoden der Siboga-Expedi-
tion.—Uitkomsten op Zod6logisch, Botanisch,
Oceanographisch en Geologisch Gebied verza-
meld in Nederlandsch Oost-Indié, 1899-1900
30:1—40. Leiden: E. J. Brill.
Poulsen, E. M. 1962. Ostracoda-Myodocopa, 1: Cy-
pridiniformes-Cypridinidae.—Dana Report 57:
1—414. Copenhagen: Carlsberg Foundation.
Sars, G. O. 1866. Oversigt af Norges marine Ostra-
coder.—Forhandlinger i Videnskabs-Selskabet I
Christiania 8:1—130. [Preprint, 1865.]
Skogsberg, T. 1920. Studies on marine ostracods, I:
Cypridinids, Halocyprids, and Polycopids.—
Zoologiska Bidrag fran Uppsala (supplement
1): 1-784.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):480—485. 2000.
A new record of Cornechiniscus madagascariensis Maucci, 1993
(Tardigrada: Echiniscidae) from India
Wataru Abe and Masatsune Takeda
(WA, MT) Department of Biological Sciences, Graduate School of Science,
The University of Tokyo;
(MT) Department of Zoology, National Science Museum, 3-23-1 Hyakunincho, Shinjuku-ku,
Tokyo, 169-0073 Japan
Abstract.—Cornechiniscus madagascariensis Maucci, 1993 which has been
known only from its type locality, Madagascar, is reported from Himachal
Pradesh, northern India. Some taxonomic emendations to the original descrip-
tion are made from the paratypes.
Recently, through the courtesy of Dr. H.
Dastych of the Zoologisches Institut und
Zoologisches Museum der Universitat
Hamburg, we had an opportunity to ex-
amine an unidentified Cornechiniscus spe-
cies collected from Himachal Pradesh,
northern India. After the direct comparison
with the paratypes of C. madagascariensis
Maucci, 1993 deposited in the Museo Civ-
ico di Storia Naturale di Verona, it was
concluded that the specimen from India
should be identified with C. madagascar-
iensis Which has previously been known
only from its type locality, Madagascar. In
this paper, we describe this species in de-
tail based on the specimen from India, and
make some taxonomic emendations from
the paratypes.
The specimens mounted on the micro-
slides were closely examined by a phase
and Nomarski differential interference con-
trast microscope (Zeiss Axiophot) at maxi-
mum magnification (X2500), illustrated
with the aid of camera lucida, and measured
using an eyepiece micrometer.
Terminology is mainly that used in Abe
et al. (1998). Abbreviations used in the text
are as follows, CT: Museo Civico di Storia
Naturale di Verona (Italy), ZMH: Zoolo-
gisches Institut und Zoologisches Museum
der Universitat Hamburg (Germany).
Genus Cornechiniscus Maucci &
Ramazzotti, 1981
Diagnosis.—Echiniscidae with horn-
shaped cirrus A. Granulation on body sur-
face consists of cuticular swellings. Pseu-
dosegmental plate present. Secondary clava
hemispherical. Venter with longitudinal cu-
ticular grooves.
Type species.—Echiniscus cornutus Ri-
chters, 1907
Cornechiniscus madagascariensis Maucci,
1993
Figs. 1, 2, 3A-E
Cornechiniscus madagascariensis Maucci,
1993:383, figs. 1—4.—Antananarivo,
Madagascar.
Material examined.—Paratypes: two
adult females, Madagascar ‘‘Antsirabe, 8/v/
1989, Cornechiniscus lobatus madagascar-
iensis Maucci, paratipo, 19/v’’ mounted in
polyvinyl-lactophenol (CT 13915, 13918).
One adult female, India ‘“‘Himal [sic Hi-
machal] Pradesh, 1500 m asl., moss from
rocks, Sep 1976, lg. J. Btoszyk, (T. 4), Cor-
nechiniscus”’ (ZMH).
Comparative material.—Cornechiniscus
lobatus (Ramazzotti, 1943): syntypes: Italy
**T-44, Pseudechiniscus cornutus f. lobata,
Montirone (Abano), 12-5-42, G. Ramazzot-
ti, 1 Muta con 2 uova, 6 [sic 7] Individui,
VOLUME 113, NUMBER 2
forma lobata, (Muschi su roccia), -solo
Faure-, 24, TIPO, P. cornutus f. lobata’”’
(CP):
Female from India.—Body length 295.0
wm excluding leg IV, width 136.0 wm; body
width : body length, 1:2.17. Eyespot well-
marked, black, subelliptical, 9.7 4m wide,
situated in posterior part of head plate.
Body color evenly translucent in preserved
material.
Dorsal plates thick. Dorsal surface in-
cluding intersegmental lateral plates wholly
covered with fairly coarse granulation that
consists of pillar-shaped cuticular granules;
adjacent granules distantly spaced, inter-
connected with conspicuous cuticular stri-
ae; granules near edge of each plate much
smaller than those at median part; granules
on scapular, segmental paired, median, in-
tersegmental lateral (1 and 2), pseudoseg-
mental, and terminal plates ca. 2.5 wm in
diameter; granules on head plate and ante-
rior part of neck plate ca. 1.5 wm in di-
ameter; granules on posterior part of neck
plate ca. 0.2 um in diameter. Lateral region
of neck plate and basis for cirrus A only
with very fine, densely distributed puncta-
tions that consist of pillar structures of epi-
cuticle.
Head plate well-developed, with two zig-
zag-sutures, showing facetted appearance.
Neck plate conspicuous, subdivided ver-
tically into three parts; posterior part of
neck plate developed as a lobe which is nar-
rowly overlapping anteriormost part of
scapular plate.
Scapular plate well-developed, anterior
with poorly defined W-shaped sculpture;
posterior to W-shaped sculpture, scapular
plate shallowly subdivided vertically at
middorsal line of body; an oblique cuticular
line at each lateral side.
Median plate 1 clearly subdivided into
anterior and posterior parts; anterior part
fairly larger than posterior part; anterior and
posterior parts trapezoidal and inverted sub-
triangular in shape, respectively.
Intersegmental lateral plates 1 and 2 sim-
ilar to each other in size and shape.
481
Segmental paired plates II and III well
developed, similar to each other in size and
shape, both with lateral segmental plates;
small, triangular spines developed posterior
part of lateral segmental plates II and III
each (spines C and D).
Median plate 2 clearly subdivided into
anterior and posterior parts; anterior part
subpentagonal, fairly larger than posterior
part; posterior edge of posterior part sinu-
ate.
Median plate 3 well-developed, rhom-
boidal, undivided.
Pseudosegmental plate clearly subdivid-
ed vertically at middorsal line of body; lobe
on its posterior margin well-developed, un-
paired, but shallowly bilobate, 11.4 pm
high, 33 wm wide at base; tips of lobe
smooth, without spine.
Terminal plate with obvious, long inci-
sion which is slightly effaced near middor-
sal line of body; minuscule, triangular spine
developed posteriormost of incision (spine
BE).
Leg plates developed on outer surfaces
of legs I-III and dorsal surface of leg IV;
each leg plate with coarse granulation sim-
ilar to those on dorsal plates, i.e. adjacent
granules interconnected with cuticular stri-
ae; granules larger but sparse on median
part of leg plate, smaller but dense on pe-
ripheral part. Plate of leg IV without true
dentate collar, but with very blunt triangular
process posterior to leg plate. Sensory or-
gan on leg I conical, 6.3 um long, 1.7 wm
wide at base; sensory organ on leg IV pa-
pillate, 6.9 um long, 5.1 wm wide at base.
Dense patches of cuticular granulation
situated below mouth opening, between
each pair of legs I-III, and around gono-
pore; these patches each without perceptible
edge. Venter longitudinally costate, with
some shallow, linear grooves that devel-
oped from cuticular patch between leg I to
gonopore, along with midline of body.
Mouth opening and cephalic sensory or-
gans situated ventrally. Internal cirrus on-
ion-shaped, tapering sharply toward tip, 8.6
wm long, 5.1 4m wide at base; external cir-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs ©.
view; C, cirrus A and primary clava on the left, dorsal view; D, sensory organ I; E, sensory organ IV; E left
side of segmental paired plate III and pseudosegmental plate; G, external claw of leg I; H, internal claw of leg
IV. (A-E, G, H: ZMH; adult female; Himachal Pradesh, N India. F: CT 13918; paratype adult female; Antsirabe,
Madagascar).
rus stout, tapering gradually, 21.4 wm long,
4.0 wm wide at base; both internal and ex-
ternal cirri without true cirrophores, but
their proximal portions swollen in appear-
ance; furthermore, internal and external cir-
ri each with hemispherical cuticular swell-
ing beneath own base; both sides of cutic-
Cornechiniscus madagascariensis Maucci, 1993. A, habitus, dorsal view; B, cephalic region, ventral
ular swellings of external cirri interconnect
each other with arcuate cuticular line. Sec-
ondary clava hemispherical, 7.4 wm in di-
ameter, situated nearer to internal cirrus
than to external cirrus.
Cirrus A clearly longer than external cir-
rus, 28.6 wm long, 9.7% of body length, 6.9
VOLUME 113, NUMBER 2
Aad
ae +)
.
>
*
Fig. 2. Cornechiniscus madagascariensis Maucci,
1993. Habitus, dorsal view. (ZMH; adult female; Hi-
machal Pradesh, N India). Differential interference
contrast. Scale = 50 wm.
14m wide at base, directed laterad; cirrus A
provided with normally developed cirro-
phore; primary clava situated just behind
cirrophore of cirrus A, clavate, curved,
apex directed posteriad, 8.0 wm long, 4.8
14m in diameter at base.
Claws I-III distally curved, whereas claw
IV less curved compared with claws I-III;
all claws thickened basally, without spur;
internal claw slightly longer than external
claw on all legs; claws I-III 12.8-14.3 wm
long; claw IV clearly longer than claws I-
Ill, ca. 18 wm long.
Female gonopore normal, consists of ro-
sette-like structure, 16 wm in diameter.
Anus large, situated posteriad, near base of
legs IV.
483
Emendatory notes on the paratypes.—
Body 282.1 and 322.4 wm long. Neck plate
tripartite vertically. Scapular plate orna-
mented with ill-defined W-shaped sculp-
ture. Cuticular grooves on venter well-
marked. Small spines developed at posi-
tions C, D, and E. Other characters in con-
cordance with Maucci (1993).
Remarks.—We examined two paratypes
(engths 295.0 and 322.4 wm) of C. mada-
gascariensis. The specimen from India is
almost identical with the paratypes includ-
ing the following important characters: pat-
tern of the dorsal granulation, shape and
size of each claw, morphology of the ce-
phalic and leg sensory organs. Although we
were able to examine only one specimen
from India, there seems to be no problem
for us to identify the specimen with C. mad-
agascariensis.
Maucci (1993) considered that C. mad-
agascariensis is most closely related to C.
lobatus Ramazzotti, 1943, and we agree
with him on this point. He pointed out that
the consistent absence of the dorsal and lat-
eral body spines is one of the most impor-
tant characters to distinguish C. madagas-
cariensis from C. lobatus. We confirmed,
however, that, as in C. lobatus, the small
spines are actually present at positions C,
D, and E also in C. madagascariensis (both
in the paratypes and specimen from India).
There are some reports of C. lobatus pro-
vided with cuticular striae among dorsal
granules, viz. Binda & Pilato (1972) from
Sicily (Italy), Dastych (1979) from Afghan-
istan, and Moon & Kim (1991) from South
Korea. Furthermore, we have verified re-
cently that the cuticular striae are rather
conspicuous, especially in the terminal
plate, in the syntypes of C. lobatus (Fig.
3F). The cuticular striae were not men-
tioned in the original description (Ramaz-
zotti, 1943) and subsequent redescription
(Maucci, 1979).
From the evidence discussed above, the
presence or absence of body spines and cu-
ticular striae among granules cannot be
used as taxonomic characters in discrimi-
484 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. A-E: Cornechiniscus madagascariensis Maucci, 1993. A, anterior body, dorsal view; B, pseudo-
segmental and terminal plates, dorsal view; C, posterior body, dorsal view; D, venter between legs I to III,
showing cuticular grooves; E, mouth and cephalic sensory organs. (A, B, E: ZMH; adult female; Himachal
Pradesh, N India. C, D: CT 13918; paratype adult female; Antsirabe, Madagascar). F: C. lobatus (Ramazzotti,
1943), pseudosegmental and terminal plates, dorsal view (CT; a syntype adult female). A, B, D—F: differential
interference contrast; C: phase contrast. Scales = 20 wm (A-D, F), 8 wm (E).
nating C. madagascariensis from C. loba- important criterion to distinguish C. mada-
tus. gascariensis from C. lobatus (Fig. 3F).
In C. madagascariensis, however, gran-
ulation on body surface consists of fairly Discussion
large, distantly spaced granules (Fig. 3A— In the original description of C. mada-
C), and this can be considered as the most gascariensis, Maucci (1993) implied that
VOLUME 113, NUMBER 2
this species may be referred to a subspecies
of a cosmopolitan species, C. lobatus Ra-
mazzotti, 1943. This also can be inferred
from his specimen labels on microslides
(see Material examined). Considering the
present record from India which is quite far
from Madagascar, however, the subspecific
status cannot be supported. We are in agree-
ment with Maucci (1993) who recognized
C. madagascariensis as a good species
based on the differences mainly discussed
above.
We have confirmed that the ventral cutic-
ular grooves, which have been omitted from
the previous descriptions, are also common-
ly developed in many other Cornechiniscus
species (unpublished data). In Pseudechin-
iscus, which is a closely related genus to
Cornechiniscus, the venter is ornamented
with net-like pattern in many species (Das-
tych 1984, Kendall-Fite & Nelson 1996). It
is thus mentioned at present that the lon-
gitudinal groove character-condition is
unique for the genus Cornechiniscus.
Acknowledgments
We thank Dr. Hieronymus Dastych of the
Zoologisches Institut und Zoologisches
Museum der Universitat Hamburg for loan
of the valuable specimen and comments on
the manuscript, to Prof. Roberto Bertolani
of the Dipartimento di Biologia Animale,
Universita di Modena for the arrangements
of loan of the paratypes, and to Prof. Clark
W. Beasely of the McMurry University and
485
Prof. Diane R. Nelson of the East Tennes-
see State University for comments on the
manuscript.
Literature Cited
Abe W., K. Utsugi, & M. Takeda. 1998. Pseudechin-
iscus asper, a new Tardigrada (Heterotardigra-
da: Echiniscidae) from Hokkaido, northern Ja-
pan.—Proceedings of the Biological Society of
Washington 111:843—848.
Binda, M. G., & G. Pilato. 1972. Tardigradi muscicoli
di Sicilia. (IV Nota).—Bollettino delle Sedute
della Accademia Gioenia di Scienze Naturali in
Catania, Serie IV 11:47—60.
Dastych, H. 1979. Tardigrada from Afghanistan with
a description of Pseudechiniscus schrammi, sp.
nov.—Bulletin de la Société des Amis des Sci-
ences et des Lettres de Poznan, Série D: Sci-
ences Biologiques 19:99—108.
. 1984. The Tardigrada from Antarctic with de-
scription of several new species.—Acta Zoolo-
gica Cracoviensia 27:377—436.
Kendall-Fite, K., & D. R. Nelson. 1996. Two new spe-
cies of tardigrades from Short Mountain, Ten-
nessee, USA.—Zoological Journal of the Lin-
nean Society 116:205—214.
Maucci, W. 1979. I Pseudechiniscus del gruppo cor-
nutus, con descrizione di una nuova specie (Tar-
digrada, Echiniscidae).—Zeszyty Naukowe Un-
iwersytetu Jagiellonskiego, Prace Zoologiczne
25:107-124.
. 1993 [dated 1990]. Prime notizie su tardigradi
«terrestri» del Madagascar con descrizione di
tre specie nuove.—Bollettino del Museo Civico
di Storia Naturale di Verona 17:381—391.
Moon, S. Y., & W. Kim. 1991. Systematic study on
the tardigrades from Korea: new records of five
tardigrade species from Korea.—Korean Jour-
nal of Systematic Zoology 7:225—232.
Ramazzotti, G. 1943. Nuova varieta del tardigrado
Pseudechiniscus cornutus.—Natura (Milano)
34:89—90.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):486—492. 2000.
Ophryotrocha lipscombae, a new species and a possible connection
between ctenognath and labidognath-prionognath eunicean
worms (Polychaeta)
Hua Lu and Kristian Fauchald
(HL)Department of Biological Sciences, the George Washington University,
Washington, DC 20052, U.S.A.;
(HL, KF) Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, DC 20560-0163, U.S.A.
Abstract.—One new species of dorvilleid, Ophryotrocha lipscombae, is de-
scribed based on material from the U.S. Atlantic slope. This new species is
unique in its presence of 5 pairs of accessory plates, which are never found in
any other dorvilleids, but are present in all labidognath-prionognath euniceans.
A brief discussion of the phylogenetic position of Ophryotrocha lipscombae in
the Eunicida is provided.
Ophryotrocha has been extensively stud-
ied by many polychaetologists (e.g., Jumars
1974, Akesson 1984, Hilbig & Blake 1991,
Eibye-Jacobsen & Kristensen 1994, Pleijel
& Eide 1996); this may due to a number of
factors: Ophryotrocha is common in both
shallow and deep water environments; fur-
thermore, it is easily cultured for study of
reproductive and developmental patterns; in
addition, its phylogenetic position within the
Dorvilleidae has turned out to be interesting.
As a part of the study of Eunicida phylog-
eny, a new Ophryotrocha species was found
by chance. In a jar containing mixed small-
sized eunicean specimens labeled only as
‘‘Runicidae’’, not only one eunicid larva was
found, but also two species of Ophryotrocha
and one juvenile lumbrinerid could be iden-
tified. All specimens were observed using
stereo and compound light microscopes; il-
lustrations were made using a camera lucida.
Abbreviation.—Mx refers to maxillae.
The species description is in a similar for-
mat as that used by Hilbig & Blake (1991).
Ophryotrocha lipscombae, new species
Figs. 1-4.
Material examined.—North Atlantic
Ocean, United States, 110 miles south of
Woods Hole, Massachusetts, 29 Jul 1977,
39°47'N, 70°40’W, Alvin Dive 773, STA
DOS-1 (N-34), 1830 m (Holotype, USNM
186571; 7 paratypes, USNM 186572).
North Atlantic Ocean, United States, 110
miles south of Woods Hole, Massachusetts,
15 Jun 1976, 39°47'N, 70°40’'W, Alvin Dive
658, 1830 m (Paratype, USNM 186573).
Description.—Holotype complete with
31 chaetigers, 2.75 mm long, 0.41 mm wide
(Fig. 1A). Other complete specimens 0.90—
2.43 mm long, 0.17—0.44 mm wide (Figs.
1B, C; 2). Largest specimen incomplete
with width of 0.77 mm. Body slender, oval
in cross section. Chaetigers two times wider
than long throughout body. One ciliary gir-
dle present on each peristomial ring and
each chaetiger. Color whitish in specimen
collected in 1976 and brownish in speci-
mens collected in 1977.
Prostomium distally bluntly triangular in
smaller specimens (Fig. 1B), rounded in
larger ones (Fig. 1A); two times longer than
wide in holotype. Two short and stout,
knob-like antennae, not reaching anterior
end of prostomium. No evidence of palps
and eyes observed in any specimen. Prosto-
mium same length and width as peristom-
ium. Peristomium consisting of two apo-
VOLUME 113, NUMBER 2
dous, achaetigerous rings, each ring slightly
shorter than adjacent chaetigers. Distinct
lateral incisions present between peristo-
mial rings; first peristomial ring distinctly
fused with prostomium.
Parapodia uniramous, with one acicular
lobe; inferiormost simple chaetae in a sep-
arate chaetal lobe. Dorsal and ventral cirri
absent. Chaetae of three types: type 1, 3—4
serrated cultriform simple chaetae (Fig. 1E)
tapering to slender distal teeth in supra-
acicular fascicle (Fig. 1D), their subdistal
end finely serrated; type 2, 4—7 (the number
variable in different body regions and dif-
ferent specimens) heterogomph bifid com-
pound falcigers (Fig. 1F) arranged in two
rows in subacicular fascicle, their blades
short and serrated; type 3, single, slender,
simple cultriform chaeta (Fig. 1G) emerg-
ing from long chaetal lobe in inferiormost
position, forming an angle with acicula at
the base. Each parapodium with one sharply
pointed acicula distinctly deeper into para-
podium than other chaetae. Pygidium rela-
tively long, as long as last two chaetigers
in 24-chaetiger specimen. Anal cirri easily '
broken, one small knob-like pair observed
on 24-chaetiger specimen. No distinct mid-
dle pygidial stylus present.
Mandible rod-like, black, two pieces
forming X-shape. In 0.17 mm-wide speci-
men, distal end of each mandible piece with
about 20 small teeth (Fig. 3B); in 0.44 mm-
wide specimen, distal end of mandible
without teeth (Fig. 3C). Maxillae K-type,
with 8 paired pieces in roughly four rows;
on each side, MX-I, II, and V-VII each
forming one main row, while Mx-III and
Mx-IV each as a separate row sitting out-
side (Fig. 4A). Mx-I (Figs. 3A, 4A) heavy,
generally ice-tong shaped and facing each
other, with 3 large teeth in addition to main
fang. Mx-II (Figs. 3A, 4A) thin plates,
forming an arc over distal part of Mx-l.
Mx-II with about 12 large teeth and a few
irregularly placed small teeth in the middle.
Mx-V to Mx-VIII (Fig. 4A) smaller than
Mx-lIl, but structurally similar, plate shaped,
with 4—8 large teeth and several small teeth.
487
Mx-III and Mx-IV elongate and fang-
shaped with additional 1 or 2 smaller teeth,
sitting anterior to Mx-II and outside of Mx-
V. One pair of short carriers (Figs. 3A, 4A)
fused to each other, and to posterior end of
Mx-I. Two rows of 5 sclerotinized black ac-
cessory plates (Fig. 4A) in erect position
inside main maxilla row, each plate corre-
sponds to Mx-II and Mx-V to VIII. Jaw
structure symmetrical.
Remarks.—Ophryotrocha lipscombae
can be easily distinguished from other
Ophryotrocha species by the presence of 3
teeth in addition to the main fang on Mx-l,
the presence of five accessory plates and
the anterior end formed by the prostomium
and the first peristomial ring. The above
features are novel in dorvilleids, especially
the accessory plates which have never been
reported in the Dorvilleidae but are present
in all other major families of the Eunicida
(personal observation). When it is com-
pared to other dorvilleids, such as O. akes-
soni and O. geryonicola, O. lipscombae ap-
pears to have a larger jaw apparatus for a
similarly sized specimen; its maxillae can
reach through the anterior four to five chae-
tigers, while those of the other species usu-
ally reach through the anterior two chaetig-
ers. The over-all jaw structure is similar in
all four Ophryotrocha lipscombae_ speci-
mens dissected whose size ranged from
0.17 mm to 0.77 mm wide.
This species reaches a relatively large
size (as wide as 0.77 mm) among dorvil-
leids. While no gametes have been ob-
served, we do not believe these specimens
to be juveniles of any other reported euni-
cean worms. The type material is deposited
in the National Museum of Natural History,
Smithsonian Institution, Washington D.C.
Geographic distribution.—110 miles
south of Woods Hole, Massachusetts, At-
lantic slope.
Etymology.—The species is named after
Prof. Diana Lipscomb of George Washing-
ton University for her many contributions
to systematic biology.
488 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
fn
EPR G
Fig. 1. Ophryotrocha lipscombae: A, anterior end, 31-chaetiger specimen, dorsal view; B, anterior end, 18-
chaetiger specimen, ventral view; C, posterior end, 24-chaetiger specimen, ventral view; D, parapodium 3,
anterior view; E, supra-acicular chaetae; K compound falciger; G, most inferior simple chaeta.
VOLUME 113, NUMBER 2
3.0
2.5
= NS
on ro)
Body size (mm)
=
0.5
0.0
16 18 20 22
Fig. 2.
Discussion
The eunicean worms share a set of com-
plex jaws composed of a ventral mandible
and dorsal maxillae, and they are classified
into five types based on jaw morphology:
labidognath (Ehlers 1868) includes Eunici-
dae, Onuphidae, Lumbrineridae and Hart-
maniellidae. This type has a pair of well
separated short carriers; Oenonidae have
prionognath jaws (Colbath 1989, Fauchald
& Rouse 1997) characterized by the pres-
ence of a median plate inside the muscular
bulb; Dorvilleidae has 4 rows of maxillae
and is called ctenognath; xenognath (Mier-
zejewski & Mierzejewska 1975) and pla-
cognath (Kielan-Jaworowska 1966) are rep-
resented only by fossil taxa. The above
classification of eunicean jaws might be ar-
bitrary if cladistic tree-thinking is applied.
489
30
Chaetiger number of complete specimen
24 26 28 32
Correlation between body size and chaetiger number in Ophryotrocha lipscombae.
Usually ctenognath is considered as mono-
phyletic, and it is a sister group of labidog-
nath-prionognath taxa (Kielan-Jaworowska
1966, Kozur 1970, Jumars 1974, Orensanz
1990). The assessment of jaw homology
can be relatively easily made between
prionognath and labidognath (Orensanz
1990), but both are generally considered
difficult to compare to the ctenognath. The
overall phylogenetic construction of Euni-
cida by Tzetlin (1980) may be problematic
when a cladistic analysis is performed
based on additional evidence (unpublished
data); however, his scheme connecting the
K-type Ophryotrocha jaw with those of la-
bidognath-prionognath jaws is reasonable
based on the following evidence: first, the
Mx-I of labidognath-prionognath is homol-
ogous with the Mx-I of both P- and K-type
490
0.05mm
0.05mm
0.1mm
rien 2:
Se
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
a
Ophryotrocha lipscombae: A. jaw structure in dorsal view, only Mx-I, Mx-II and the carrier are
depicted; B. mandible of 18-chaetiger specimen, dorsal view; C. mandible of 31-chaetiger specimen, dorsal view.
Ophryotrocha, based on extensive studies
of larval and adult jaw morphology of Dor-
villeidae, Onuphidae and Oenonidae (H. L.,
unpublished data); second, the accessory
plates, or attachment lamellae (Paxton
1986), are present in all members of the
labidognath-prionognath taxon (personal
observation), and the presence of such
plates in Ophryotrocha lipscombae is the
first record for any dorvilleid. Accessory
plate numbered 5 pairs in O. lipscombae, 3
pairs in Oenonidae (Fig. 4B) and Lumbri-
neridae (Fig. 4C), and 2—3 in Onuphidae
and Eunicidae. All accessory plates are lo-
cated median to the corresponding maxillae
in erect position; no plate corresponds to
Mx-lI.
The presence of accessory plates may
provide a substantial primary homology for
O. lipscombae and the labidognath-prion-
ognath taxa; the relationship will be tested
in a systematic study of Eunicida based on
morphology, ontogeny and fossil data. At
present, the new species is considered as a
member of Ophryotrocha based on its over-
all morphological characters. Though the
phylogenies of Ophryotrocha (Pleyel &
Eide 1996) and Dorvilleidae (Jumars 1974,
Westheide 1982, Hilbig & Blake 1991, Ei-
bye-Jacobsen & Kristensen 1994) have
been studied using various methods, better
overall understanding might be gained from
a relatively broader study of the phylogeny
of the order Eunicida, including detailed
studies of members of all major groups.
Acknowledgments
The authors would like to thank Linda A.
Ward, Cheryl Bright, William Moser, and
VOLUME 113, NUMBER 2
491
Fig. 4. A. Detailed jaw structure of 31-chaetiger specimen, Ophryotrocha lipscombae, in ventral view; note
that left and right sides are cut apart, the carrier is broken, and the whole structure is displayed in an unnatural
way in order to show the detailed structure; B. jaw structure in ventral view, Arabella iricolor (USNM 10355);
C. jaw structure in ventral view, Lumbrineris zonata (USNM 30611).
other colleagues from the Department of In-
vertebrate Zoology, NMNH, Smithsonian
Institution, and Diana Lipscomb and the
Systematics Group, Department of Biolog-
ical Sciences, the George Washington Uni-
versity. We thank Stephen Gardiner and
Kirk Fitzhugh for their reviews. HL is sup-
ported by the Research Enhancement Funds
and Weintraub Research Fellowship from
the George Washington University, and re-
492
search contract 9822uu04278 from the
Smithsonian Institution.
Literature Cited
Akesson, B. 1984. Speciation in genus Ophryotrocha.
Pp. 299-316 in A. Fischer and H. D. Pfan-
nenestiel, eds., Polychaete reproduction, pro-
gress in comparative reproductive biology, vol.
29, Fortschritte der Zoologie (Stuttgart: Gustav
Fischer Verlag).
Colbath, G. K. 1989. Revision of the family Lysare-
tidae, and recognition of the family Oenonidae
Kinberg, 1865 (Eunicida; Polychaeta).—Pro-
ceedings of the Biological Society of Washing-
ton 102:116—123.
Ehlers, E. 1868. Die Borstenwiirmer (Annelida: Chae-
topoda) nach systematischen und anatomischen
Untereuchungen dargestellt, W. Engelmann,
Leipzig, 748 pp.
Eibye-Jacobsen, D., & R. M. Kristensen. 1994. A new
genus and species of Dorvilleidae (Annelida,
Polychaeta) from Bermuda, with a phylogenetic
analysis of Dorvilleidae, Iphitimidae and Di-
nophilidae.—Zoologica Scripta 23:107—131.
Fauchald, K., & G. Rouse 1997. Polychaeta system-
atics: past and present.—Zoologica scripta
26(2):71-138.
Hilbig, B., & J. Blake 1991. Dorvilleid (Annelida:
Polychaeta) from the U.S. Atlantic slope and
rise. Description of two new genera and 14 new
species, with a generic revision of Ophryotro-
cha.—Zoologica Scripta 20 (2):147-183.
Jumars, P. 1974. A generic revision of the Dorvilleidae
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
(Polychaeta), with six new species form the
deep North Pacific.—Zoological Journal of the
Linnean Society 54(2):101—135.
Kielan-Jaworowska, S. 1966. Polychaete jaw appara-
tuses from the Ordovician and Silurian of Po-
land and a comparison with modern forms.—
Palaeontologia Polonica 16:1—152.
Kozur, H. 1970. Zur Klassifikation und phylogene-
tischen Entwicklung der fossilen Phyllodocida
und Eunicida (Polychaeta).—Freiberger For-
schungshefte 260 C:35-81.
Mierzejewski, P., & G. Mierzejewski 1975. Xenognath
type of polychaete jaw apparatuses.—Acta Pa-
laeontologica Polonica 20:437—443.
Orensanz, J. M. 1990. The Eunicemorph polychaete
annelids from Antarctic and Subantarctic seas.
With addenda to the Eunicemorpha of Argen-
tina, Chile, New Zealand, Australia, and the
southern Indian Ocean.—Antarctic Research
Series 21:1—183.
Paxton, H. 1986. Generic revision and relationships of
the family Onuphidae (Annelida: Polychae-
ta).—Records of the Australian Museum 38:1—
74.
Pleijel, EF, & R. Eide 1996. The phylogeny of Ophry-
otrocha (Dorvilleidae: Eunicida: Polychaeta).—
Journal of Natural History 30:647—659.
Tzetlin, A. B. 1980. Ophryotrocha schubravyi sp. n.
and the problem of evolution of the mouth parts
in the Eunicemorpha (Polychaeta).—Zoologi-
cheskii Zhurnal 59:66—676.
Westheide, E. 1982. /kosipodus carolinensis gen. et sp.
new., an interstitial neotenic polychaete from
North Carolina U.S.A., and its phylogenetic re-
lationships within Dorvilleidae.—Zoological
Scripta 11:117—126.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):493—499. 2000.
Aphrodita bisetosa (Polychaeta: Aphroditidae), a new species of sea
mouse from the southeastern Pacific Ocean off central Chile
Nicolas Rozbaczylo and Elba Canahuire
(NR) Departamento de Ecologia, Facultad de Ciencias Bioldégicas, P. Universidad Cat6lica de
Chile, Casilla 114-D, Santiago, Chile, e-mail: nrozbac @ genes.bio.puc.cl;
(EC) Facultad de Ciencias Biol6gicas, Universidad Nacional Mayor de San Marcos, Lima, Pert;
presently: Programa de Magister en Ciencias menci6n Zoologia, Escuela de Graduados,
Universidad de Concepcién
Abstract.—A new species of Aphroditidae, Aphrodita bisetosa from the
southeastern Pacific Ocean off central Chile, is described. Specimens were
collected in soft bottoms, 27—37 km from the coast line in front of Los Vilos
(31°56’S) and Papudo (32°31'S), at 250-400 m depth. The new species was
compared with A. magellanica Malard, 1891 from the Magellan area, Chile
and A. alta Kinberg, 1855 from Rio de Janeiro, Brazil, and the Antarctic region,
with which it appears to be more closely related.
Aphroditids are commonly named “‘sea
mice,’’ because of the thick mat of very fine
notopodial fibers which form a felt-like
covering over the dorsum, giving them a
furry appearance. Although they are rather
large and conspicuous inhabitants of marine
soft muddy bottoms, they are frequently ab-
sent from most collections because of the
great depths at which they usually live, and
in general they have been poorly studied.
The most recent and comprehensive study
of aphroditids was by Hutchings & McRae
(1993) on species found in Australian wa-
ters and the Indonesian Archipelago.
There is almost no information about
aphroditids of the southeastern Pacific
Ocean along the Chilean coast. Only one
species, Aphrodita magellanica Malard,
1891, has been previously recorded from
the Magellan area (49°S) south to Cape
Horn (56°S) in southern Chile (Rozbaczylo
1985); the specimens, had been collected
during the Challenger Expedition (1873-
1876), and identified by McIntosh (1885) as
Aphrodita echidna Quatrefages.
Four specimens of aphroditids collected
by commercial shrimp trawlers at two sites
off central Chile were given to the first au-
thor for study; after examination, they were
considered as belonging to a new species.
Materials and Methods
Specimens were collected by the shrimp-
ing boat Goden Wind, as part of a benthic
survey obtained 27—37 km from the coast
between Los Vilos and Papudo, in October
1976. At 2 of 5 stations sampled, specimens
of aphroditids were found in sandy-mud
bottom (Fig. 1). Additional information on
the accompanying macrofauna collected
during the trawls can be found in Andrade
(1986).
Polychaetes were fixed in 4% formalin
and preserved in 70% ethanol. Figures were
prepared with a drawing tube on a Wild M-
5 stereoscopic microscope and a Leitz com-
pound microscope.
Type specimens of the new species are
deposited in the National Museum of Nat-
ural History, Smithsonian Institution,
Washington, D.C. (USNM), and the Sala de
Sistematica, Departamento de Ecologfa,
Pontificia Universidad Catdélica de Chile,
Santiago (SSUC).
494
Aphrodita bisetosa, new species
Figs. 1-3
Material examined.—Central Chile: in
front of Los Vilos,. Stieilecay 3156'S,
71°49'W, 300—400 m, H. Andrade, coll., 14
Oct 1976, holotype (USNM 186512) and
female paratype (SSUC 6868); in front of
Papudo, St. 3, ca. 32°31'S, 71°47'W, 250—
280 m, H. Andrade, coll., 14 Oct 1976, par-
atype (N° 2 USNM 186513) and female
paratype (SSUC 6869).
Description.—Holotype. Body ovate,
arched dorsally, widest at setigers 14—16,
with tapering caudal region (Fig. 2a); 27
mm long, 16 mm wide, excluding setae,
with 41 setigers. Dorsum with thick felt, ap-
proximately 1.5 mm thick at middle region
of body, with fine sediment entrapped giv-
ing it greyish appearance. Ventral surface of
body whitish, covered with minute spheri-
cal papillae (Fig. 2d).
Prostomium small, rounded with pair of
ocular areas, pale to light brown in color,
located on slightly raised prominences (Fig.
2b). Median antenna with basal ceratophore
and elongated style, slightly shorter than
prostomium, attached dorsally, near anterior
border of prostomium. Palps_ biarticulate,
finely papillated, wide basally, tapering
gradually, extending approximately to sixth
setiger. Facial tubercle minutely papillated,
approximately half of length of prostomi-
um, partly hidden by palps dorsally and ex-
tending ventrally as digitiform process over
mouth (Fig. 2b, c).
Elytra (Fig. 2e, f) 15 pairs, completely
hidden by dorsal feltage, on setigers 2, 4,
5, 1, 9, sex 25, 2831; imbricate, *complete-
ly covering dorsum, semi-transparent;
smooth except for few scattered microscop-
ic digitiform papillae (Fig. 2g), mostly con-
centrated on inner lateral area of upper sur-
face and less on posterior area of elytra.
First pair of elytra smallest, gradually in-
creasing in size to approximately pair 7-9,
then decreasing posteriorly. First and sec-
ond pair of elytra ovate, longer than wide,
with elytrophores attached centrally and lat-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
erally, respectively; following elytra wider
than long; last three pairs noticeably longer
than wide; last pair smaller.
Dorsal tubercles from segment 6 to 30
provided with branched fimbriated papillae
on posterolateral margins (Fig. 3d).
First setiger or tentacular segment with
elongated, uniramous parapodia, flattened,
projecting anteriorly and laterally to prosto-
mium (Fig. 2b), with one tuft of fine silky
fibers emerging dorsally forming the dorsal
felt, and 3 fascicles of fine, faintly irides-
cent, mud-covered capillary setae; one tuft
supra-acicular and two others sub-acicular;
with few scattered papillae covering para-
podium (Fig. 2b). Parapodial dorsal and
ventral cirri with cylindrical cirrophores
and subulate styles. Dorsal cirri approxi-
mately one-third length of palps. Ventral
cirri slightly shorter than dorsal.
Following setigers with biramous para-
podia. Second setiger with first pair of el-
ytra. Notopodia rectangular, with one tuft of
fine silky fibers forming dorsal felt, 2 supra-
acicular fascicles of stout acicular setae,
with hooked tips, similar in shape and dis-
tribution to that of parapodium 3, and a sub-
acicular fascicle with capillary notosetae
forming lateral fringe, and on posterior sur-
face of notopodia at level of acicula, a small
tuft of fine silky fibers forming dorsal felt.
Neuropodia cylindrical, covered with spher-
ical papillae; tip of acicula emerging at dis-
tal end of neuropodial lobe. Neurosetae,
brown, arranged in 3 tiers: upper consisting
of 2—3 acicular setae, middle 1—2 acicular
setae, and lower of numerous capillary se-
tae (Fig. 31), slender bipinnate and spirally
twisted, with two rows of thick teeth that
continue in subdistal region as fine spines,
and with distal end smooth. Ventral cirri su-
bulate, located near base of neuropodium,
about one and one-half length of neuropo-
dia (Fig. 2d).
Third setiger similar to second, but with
dorsal cirri instead of elytra. Dorsal cirri
long, subulate, approximately 3-4 times
length of ventral cirri.
Parapodia of middle region of body (Fig.
VOLUME 113, NUMBER 2
2
<
LL]
O
O
Oo
LL.
O
<x
0.
Fig, 1.
3a, b) with notopodial lobe large, nearly tri-
angular, with dorsal cirri with cirrophores
large, basally bulbous, projecting on pos-
terior faces of notopodia; styles subulate,
long, slender, smooth, directed dorsally;
dorsal cirri approximately 3—4 times longer
than ventral cirri; notoacicula stout, light
brown, emerging at vertex of distal end of
notopodial lobe.
Notopodia with tufts of fine silky fibers
emerging dorsally forming dorsal felt, with
495
LOS VILOS
PAPUDO
VALPARAISO
Map showing the stations (*) where specimens of Aphrodita bisetosa, new species, were found.
hooked ends (Fig. 31); arranged in three
main groups on cirrigerous segments, one
above and one below upper fascicle of su-
pra-acicular notosetae, and third one on
posterior surface of notopodia at level of
acicula; on elytrigerous segments appearing
in two main groups, one between upper and
lower fascicle of long supra-acicular noto-
setae, and one on posterior surface of no-
topodia at level of acicula. Three fascicles
of notosetae present, one sub-acicular and
496 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
i, @ees @\\\
\ AST
NC |i
|
|
Fig. 2. Aphrodita bisetosa, new species (Holotype USNM 186512). a, dorsal view, whole animal; b, dorsal
view of prostomium and first segment, with first pair of elytra omitted (right palp and dorsal parapodial cirri
missing); c, facial tubercle (Paratype SSUC 6869), antero-lateral view; d, anterior end (Paratype SSUC 6869),
ventral view; e, anterior end (Paratype USNM 186513), dorsal view, with proboscis everted and feltage set
aside, showing first three pairs of elytra, style of median antenna missing; f, ninth left elytron, inner surface
(Paratype USNM 186513); g, papillae from ninth elytron; h, posterior segments around the anus (Paratype SSUC
6869), dorsal view. Scales = 10 mm for a; 5 mm for d, e, f; | mm for b, h; 0.05 mm for g.
VOLUME 113, NUMBER 2 497
eg fle
Fig. 3. Aphrodita bisetosa, new species, a, parapodium 14 (Paratype SSUC 6868), in anterior view (tufts of
silky fibers are shown cut off); b, same parapodium in posterior view (f. p. = fimbriated papilla; s. p. = segmental
papilla); c, conical papillae from notopodium of same parapodium; d, fimbriated papillae (Paratype SSUC 6869),
from segments 6, 8, 10, and 12, respectively; e, capillary notoseta from parapodium 14; f, g, acicular notosetae
of stout type, with hooked tips, from parapodium 14; h, acicular notoseta of stouter type, with straight tip, from
parapodium 18; i, fine silky fiber, from parapodium 20; j, acicular neuroseta, with heavily bearded end, from
lower tier, parapodium 22; k, acicular neuroseta, partially bearded, from lower tier, parapodium 14; 1, capillary
bipinnate neuroseta, spirally twisted, from lower tier, parapodium 2; m, capillary, partially bipinnate neuroseta,
from lower tier, parapodium 29. Scales = 5 mm for a, b, j; | mm for d, h; 0.5 mm for c; 0.01 mm for k, 1, m;
0.005 mm for e, f, g, 1.
498
two supra-acicular. Sub-acicular fascicle
made up of numerous iridescent capillary
setae (Fig. 3e), extending laterally, with dis-
tal end straight and pointed; most covered
with fine silky fibers and fine mud resem-
bling cotton in appearance. Two supraci-
cular fascicles made up of 2 kinds of acic-
ular setae emerging through dorsal feltage:
many stout acicular setae with hooked tips
(Fig. 3f, g) and 1—2 stouter acicular setae
with straight tips (Fig. 3h), short, conical,
dark brown to blackish, spine-like, extend-
ing dorso-posteriorly (tips may be broken
off), with fine silky fibers at base. In upper
fascicle, stout protective notosetae present
(Fig. 2a), long, brown-colored basally and
shiny golden light brown color distally, ta-
pering gradually, with flexible hooked tips
(may be broken off), sometimes partially
covered with fine silky fibers. In lower fas-
cicle, stout protective notosetae short with,
distal region having fine silky fibers and
covered with fine mud presenting finger-
like appearance, peppered with ferruginous
color.
Notopodia covered with two kinds of pa-
pillae: globular, small, few, scattered on su-
bacicular area near distal margin of noto-
podial lobe, and conical (Fig. 3c), larger
and more abundant, scattered on basal area
of notopodial lobe.
Neuropodia (Fig. 3b), cylindrical, ending
distally in three step-like lobes, covered
with minute spherical papillae. Ventral cirri
small, subulate, located in middle region of
neuropodia, approximately one-third the
length of dorsal cirri.
Neurosetae stiff, stout, dark brown,
slightly curved distally, with heavily beard-
ed ends (Fig. 3j), which can be broken;
pointed subdistal tip visible through beard,
exposed when the distal tip of beard ends
lost (Fig. 3k). Neurosetae arranged in three
tiers with 2 setae in upper, 2 in middle and
4—6 in lower tier in anterior parapodium
(setiger 4), 2 in upper, 3—4 in middle and
5—9 in lower tier in middle parapodium (se-
tiger 15), and 2 in upper, 2 in middle and
7 in lower tier in parapodium of posterior
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
region of body (setiger 25). Upper group
with stoutest and longest setae; lowest most
slender and shortest. Neuroacicula, stout,
light brown, emerging through upper lobe.
Ventral neurosetae of posterior region of
body, capillaries, with two rows of teeth
subdistally (Fig. 3m), and distal region
smooth with tips from slightly curved to
straight. Neurosetae of more posterior setig-
ers all capillaries.
Anus (Fig. 2h) located dorsally, anterior
to 2 small segments. Without anal cirri.
Variation.—Additional material exam-
ined ranges from 47—80 mm long, 20-35
mm wide, excluding setae, with 42 setigers.
Body varies from ovate to elongated in
largest specimens. Fascial tubercle extends
ventrally as digitiform process over mouth
in all except one specimen (paratype SSUC
6868). Only one specimen with everted
proboscis (Fig. 2e) with opening surround-
ed by numerous, leaflike, dichotomously
branched papillae; chitinous jaws lacking.
Elytra in largest specimens are light brown
to cream in color; irregularly stained fer-
rugineous to olive in colour on inner sur-
face (Fig. 2f). Holotype without segmental
papillae, but present in largest specimens as
small knobs on posterior faces of parapodia
between rami, on segment 14, and subse-
quent setigers (Fig. 3b).
In paratypes neurosetae varied as fol-
lows: 2 setae in upper, 2—3 in middle and
3—6 in lower tier in anterior parapodium
(setiger 4), 2—3 in upper, 3—5 in middle and
10—12 in lower tier in middle parapodium
(setiger 15), and 2—3 in upper, 3—4 in mid-
dle and 7—11 in lower tier in parapodium
of posterior region of body (setiger 25).
Etymology.—The species name bisetosa
is derived from Latin bi-meaning double,
and setosa meaning with bristles, because
of the presence of two kinds of notosetae
which emerge through the dorsal feltage.
Distribution.—The species has been
found at two sites off Central Chile: between
27 and 37 km off the coast in front of Los
Vilos, 300—400 m and in front of Papudo,
250—280 m depth, in sandy-mud bottoms.
VOLUME 113, NUMBER 2
Remarks.—The new species Aphrodita
bisetosa is characterized by two kinds of
acicular setae emerging through dorsal fel-
tage: numerous stout setae with hooked
tips, and some stouter, short, dark brown,
spine-like setae, with straight tips, forming
two distinct rows of protective notosetae
along each side of body. The acicular no-
tosetae appear in two fan-shaped fascicles
of different length: an upper fascicle of long
setae, shiny golden brown-colored distally,
extending dorsomedially, nearly touching
medially, and a lower fascicle of short setae
extending backwards, bearing a mud cover,
peppered with ferruginous colour, present-
ing finger-like appearance.
Aphrodita bisetosa appears to be most
closely related to A. magellanica Malard,
1891, from the Magellan area and to A. alta
Kinberg, 1855 from Rio de Janeiro, Brazil,
and the Antarctic region. A. magellanica is
the most similar to A. bisetosa n. sp. Both
species have two rows of acicular notosetae
along each lateral region of the body that
are short, brown, and spine-like with
straight tips emerging through the dorsal
feltage. Both species have similar neurose-
tae of the acicular type, brown, with beard-
ed ends (which can be broken), and in both
species the dorsal felt is formed of very fine
silky fibers with hooked tips; the lateral re-
gions are of ferruginous color. A. bisetosa
differs from A. magellanica by having two
kinds of protective notosetae; its spine like
acicular notosetae are conical toward the
tip, 1-2 of these notosetae are present in
each supracicular tuft; ventral cirri of para-
podia lack papillae; and the body has 41—
42 setigers. In A. magellanica, in contrast,
has only one kind of protective notosetae.
The spine like notosetae are flattened to-
ward the tip, 2—3 of these notosetae can be
present in each supracicular tuft; ventral cir-
ri of parapodia possess numerous papillae
and the body has about 35 segments.
Aphrodita bisetosa resembles A. alta in
that the protective notosetae end in a hook,
the acicular neurosetae of the middle region
of the body are slightly curved and have
499
heavily bearded ends and in both species
the proboscis bears leaflike papillae sur-
rounding the opening. A. bisetosa differs
from A. alta by the following characteristics
present in the latter species: only one type
of protective notosetae that do not project
through the felting, only one type of orna-
mentation in capillary neurosetae of second
setiger; without segmental papillae and pro-
boscis with jaws.
Acknowledgments
We are greatly indebted to Dr. Héctor An-
drade who kindly made specimens available
for study. We are specially grateful to Mrs.
Clara Yanez for her skill and patience in pro-
ducing the excellent illustrations of the new
species. Two anonymous referees are thanked
for their helpful comments and suggestions
that greatly improved the manuscript.
Literature Cited
Andrade, H. 1986. Observaciones bioecoldgicas sobre
invertebrados demersales de la zona Central de
Chile. Pp. 41-56 en La Pesca en Chile. P. Ar-
ana, ed., Escuela de Ciencias del Mar, Univer-
sidad Cat6élica de Valparaiso.
Hutchings, P, & J. McRae. 1993. The Aphroditidae
(Polychaeta) from Australia, together with a re-
description of the Aphroditidae collected during
the Siboga Expedition.—Records of the Austra-
lian Museum 45(3):279—363.
Kinberg, J. G. H. 1855. Nya slagten och arter af An-
nelider.—Ofversigt af Kongliga Vetenskaps-
Akademiens Forhandlingar (Stockholm) 12:
381-388.
Malard, A. E. 1891. Sur une nouvelle aphrodite du Cap
Horn, décrite a tort par M. McIntosh comme A.
echidna (De Quatrefages).—Bulletin de la So-
ciété Philomathique de Paris Sér. 8, 3:125—127.
McIntosh, W. C. 1885. Report on the Annelida Poly-
chaeta colleted by H.M.S. Challenger during
the years 1873—1876. Jn Report on the Scientific
Results of the Voyage of H.M.S. Challenger
during the years 1873-1876 under the Com-
mand of Captain George S. Nares, R.N., ER.S.
and the Late Captain Frank Tourle Thomson,
R.N., Zoology, 12(34):1-554, pls. 1-55, and
la—39a.
Rozbaczylo, N. 1985. Los Anélidos Poliquetos de
Chile. Indice sinonimico y distribuci6n geo-
grafica de especies.—Monografias Biolégicas
3:1-284.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):500-—513. 2000.
Exogone breviantennata Hartmann-Schrdéder, 1959
(characters emended) (Annelida: Polychaeta: Syllidae), a new record
for the Bahamas with a key to selected Exogone species
Robert Zottoli and Charlene D. Long
(RZ) Fitchburg State College, Fitchburg, Massachusetts 01420, U.S.A.;
(CDL) The Buffum Group, 21 % Buffum Street, Salem, Massachusetts 01970, U.S.A.
Abstract.—The syllid polychaete Exogone breviantennata Hartmann-Schréd-
er, 1959 is reported from the Bahamas, where it was collected from epiphytic
communities on submerged red mangrove roots in Osprey Lake and Reckley
Hill Pond located on San Salvador Island. The external morphology of both
paratype and Bahamian specimens is described and species characters emended.
Observations on reproduction, biology, habitats and distribution are reported.
The essential characteristics used to identify the genus Exogone are discussed
and a key to Caribbean species with a dorsal cirrus on setiger 2 is provided.
Several polychaete species were collect-
ed from algal covered red mangrove roots
in Osprey Lake and Reckley Hill Pond,
landlocked bodies of water on San Salvador
Island in the Bahamas. One of them, re-
ported here for the first time from the Ba-
hamas, E. breviantennata WHartmann-
Schréder, 1959, has features unique to in-
terstitial polychaetes normally associated
with sand or mud. The species characters
are emended based on examination of par-
atype material and the specimens collected
from the Bahamas. Included is a discussion
of the important characters used to identify
the genus Exogone and a key to species re-
lated to E. breviantennata that are found in
the Caribbean.
Materials and Methods
Two subtidal Rhiziophora mangle prop
roots, extensively colonized by algae, were
collected from Osprey Lake, San Salvador
Island, Bahamas, 5 May 1996, and from
Osprey Lake and Reckley Hill Pond (some-
times given as Reckley Hill Settlement
Pond), 19 Jun 1997. The polychaetes were
removed, relaxed in 5% magnesium sulfate
and fixed in a 5% formalin/seawater solu-
tion. Specimens were transferred to 70%
ethanol for long-term storage. Some of the
specimens were mounted in Hoyer’s solu-
tion in order to clear body tissue so that
teeth and setae would be clearly visible
(Zottol & Long 1998) or in Glycrogel
Mountant (Gurr 1962), which allowed close
examination of body surfaces.
Measurements were made under com-
pound or dissecting microscopes with a cal-
ibrated grid. Body length was measured
from the tip of the prostomium to the end
of the pygidium; for twisted specimens, the
grid was rotated along the body. Body
width was measured from the tip of the
right neuropodium to the left across the dor-
sal surface of the fourth setiger.
Comparative material was borrowed
from the Zoologisches Museum, Hamburg
(ZMH) and the National Museum of Nat-
ural History, Smithsonian Institution
(USNM), Washington, D.C., U.S.A.
Identifying Specimens of the Genus
Exogone
Most syllids, because of their small size,
are difficult to identify, requiring careful
examination under a compound micro-
VOLUME 113, NUMBER 2
scope. For this and other reasons, Fauchald
(1977:79) stated: “‘the identification of syl-
lids [is] a time-consuming occupation.”’
Larger specimens should have the pharynx
isolated and appropriate parapodia removed
for examination under a compound micro-
scope. Smaller specimens should be cleared
in an appropriate medium such as Hoyer’s
solution (see Zottoli & Long 1998) or
mounted permanently in Glychrogel Moun-
tant (Gurr 1962). During examination, it is
important to keep in mind that what you see
may be an artifact resulting from the degree
of relaxation prior to fixation, the type of
preservative used, or the length of time
stored, especially with regard to the size
and shape of soft parts and the appearance
of setae (see Riser 1991:214). The quality
of equipment, which determines the fine-
ness of detail seen, and the experience of
the investigator, may also influence what is
observed. These factors may be responsible
for the lack of detail in many original de-
scriptions, which in some cases are now
considered incomplete because they did not
illustrate or discuss what are now consid-
ered essential characters.
The following is a discussion, on a char-
acter by character basis, of features com-
monly used to identify species of Exogone.
Size should not be used to classify speci-
mens because local and regional variations
have been inadequately studied for most
polychaetes. The degree of fusion of the
palps must be viewed both dorsally and
ventrally. The number of antennae (if not
lost) is reliable, although antenna shape,
length, and placement on the prostomium
need to be measured on a large number of
specimens; antennae are easily distorted
due to relaxation and fixation techniques.
Russell (1991) refuted the use of the num-
ber of muscle cell rows of the proventric-
ulus as a diagnostic character in the family
Syllidae, and we have found sufficient var-
iation in E. breviantennata to agree; the
number of setigers in which the proventric-
ulus is found varies according to how far
the pharynx is retracted, thus making this a
501
questionable character for species distinc-
tion. For comments regarding the useful-
ness of nuchal organs in identifying some
syllids, see Riser (1991). The shape of the
neuropodia and the shape and position of
the dorsal, ventral, and tentacular cirri are
easy to determine even in small specimens.
The presence or absence of dorsal cirri on
setiger 2 in adult specimens is an important
and stable species characteristic; however,
the apparent absence of dorsal cirri on se-
tiger 2 should be verified on several speci-
mens as the structures can easily be de-
tached. All parapodia should be scanned for
modified setae of any kind and all setae
should be carefully examined dorsally, lat-
erally, and ventrally under oil immersion
along the entire length of the animal. Setae
of the same type can appear quite different
from setiger to setiger depending on the de-
gree of rotation. For example, blades may
appear smooth if viewed from either a dor-
sal or a ventral aspect but toothed from a
lateral aspect. The tips of setal blades that
appear unidentate may, on closer examina-
tion, be seen to be bidentate. The type and
extent of denticulation or spination on the
setal shafts also become more apparent with
increasing magnification. The importance
of care in observing setae is illustrated in a
comparison of figures 75—78 in the original
description of E. breviantennata by Hart-
mann-Schroéder (1959), which show no dec-
oration on the setae, and, yet, painstaking
reexamination of a type specimen revealed
obvious decoration (blades and shafts). Fi-
nally, every specimen in a collection should
to be checked individually, to ensure that
all belong to the same species (Perkins
1981).
Genus Exogone @Orsted, 1845
Key characters.—Small worms (usually
less than 3 mm in length but up to 8 mm);
palps sometimes fused; three antennae and
one pair of tentacular cirri; dorsal and ven-
tral cirri well developed but shorter than se-
tal lobes; all appendages papilliform or
502
ovoid; eversible pharynx with single ante-
rior tooth (Fauchald 1977, Pascual et al.
1996). San Martin (1991) reviewed this ge-
nus based on his work with Cuban syllids
and expanded the number of key characters
to include presence or absence of eyes, ex-
tent of palpal fusion, presence of two long
anal cirri, smooth body surface, and repro-
ductive habits.
In summary and in decreasing order of
reliability, we use the following characters
to distinguish various species in the genus
Exogone: presence or absence of a dorsal
cirrus on setiger 2; position, shape, and
length of the antennae; shape, size, devel-
opment and placement of eyes; placement
and relative length of the proboscis and pro-
ventriculus; and shape and denticulation
and spination of setae and acicula.
Biology of the genus Exogone.—Mem-
bers of this genus have been reported from
soft and hard bottoms (Uebelacker 1984:
30-37 through 30-43), associated with
sponges (Pascual et al. 1996) and on un-
shaded algal flats dominated by Caulerpa
verticillata and Halimeda opuntia f. trilo-
bata, reported by Russell (1991). In the Ca-
nary and Madeira Islands, six species of
this genus comprised 6.6% of all poly-
chaetes collected from sponges. When they
are associated with living in sponges, some
members of this genus exhibit morpholog-
ical characters such as decrease in the blade
length of compound setae and/or blades
fused with the shafts, as is the case in other
syllid genera, e.g., Haplosyllis (Pascual et
al. 1996). However, Exogone species that
are not exclusively associated with sponges
do not show these adaptations, e.g., E. brev-
iantennata (see below). Members of the
subfamily Exogoninae usually undergo re-
productive metamorphosis that includes
changes in various external structures (eyes,
antennae, parapodial structures, and setae)
and internal structures (nephridia and mus-
culature) (Schroeder & Hermans 1975, Ku-
per & Westheide 1998). Based on the lit-
erature and our observations, E. brevian-
nenata appears to be an exception in that
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
reproductive modifications have never been
observed for either sex.
Of interest is the fact that, based on the
original descriptions of 34 species in this
genus, there is about a one to one ratio of
those that have a dorsal cirrus on setiger 2
and those that lack this character. Of the
latter group, many species were reported to
have a minute median antenna; very few of
those species were said to have a median
antenna longer than the palps. Of the for-
mer group, most species had a median an-
tenna that reached at least to the tip of the
prostomium, and many were much longer,
reaching to or beyond the palps. An excep-
tion to this is E. breviantennata, which has
a shorter median antenna, not even reaching
to the tip of the prostomium.
Exogone breviantennata Hartmann-
Schroder, 1959
characters emended
Figs. 1-5
Exogone breviantenna Hartmann-Schréder,
1959:125-127, figs. 75-78; San Martin,
1991:728-729, 730-731, fig. 8; Nunez et
al., 1992:47, fig. 3; Pascual et al., 1996:
710; 77. Aablex2:
Exogone occidentalis Westheide, 1974:113;
Russell, 1991:59—61, fig. 4 (emended).
Material examined.—El Salvador: para-
type P-14590, ZMH; Belize: Twin Cays,
West Bay, D. E. Russell, Nov 1993, Sta. M-
4, 3 specimens labelled Exogone occiden-
talis, USNM 102085; and Bahamas: San
Salvador Island (from amongst filaments of
the algae Batophora sp. and Vaucheria sp.
taken from the prop roots of Rhiziophora
mangle), Oyster Pond, R. Zottoli & C. D.
Long, 6 specimens, 4 May 1996; Osprey
Lake, R. Zottoli, 13 specimens, 5 May
1996, USNM 186554; Reckley Hill Pond,
R. Zottoh, 2 specimens, 5 May 1996 and
17 specimens, 19 Jun 1997, USNM
186555-186561; and southern edge of Bim-
ini Lagoon, from plastic mesh sponges an-
chored and floating in 1-3 feet of water,
collected at intervals up to 11 months from
VOLUME 113, NUMBER 2
1970-1971, Amy Schoener, 12 specimens
(labelled as Exogone verugera), USNM
51542, and 5 specimens, USNM 51545.
Key characters.—Syllid in the genus Ex-
ogone with dorsal cirri on all setigers and
median antenna shorter than prostomium.
Description of the paratype.—Specimen
1.95 mm long, 0.2 mm wide, 24 setigers
(Fig. 1). New segment, without setae, form-
ing between last setiger and pygidium. Pro-
stomium about four times as wide as long;
anterior and lateral margins rounded. Eyes
not discernible. In the original description
of type material, Hartmann-Schréder (1959)
described and figured four eyes, one at each
corner of a trapezoid, in the posterior half
of the prostomium. Single median and two
lateral digitiform antennae. Antennae simi-
lar in shape to tentacular, dorsal, and ventral
cirri. Palps completely fused dorsally, ex-
tend anteriorly well beyond the prostomi-
um. Tentacular segment short, about three
times as wide as long; clearly separated
from the prostomium, but not from the fol-
lowing first setiger. One pair of lateral ten-
tacular cirri, about 0.02 mm long and 0.01
mm basal width. Uniramous parapodia
(neuropodium only) as short lobes, similar
on all setigers. Digitiform dorsal (about
0.024 mm long and 0.02 mm basal width)
and ventral (about 0.024 mm long and
0.012 mm basal width) cirri. One dorsal
simple seta (3 wm basal width) per setiger;
terminates with short spines (Fig. 2A); tip
slightly concave. One heterogomph spiniger
(Fig. 2B) just below the dorsal simple seta;
blade length 20 to 30 wm, increases from
anterior to mid-body and then decreases
posteriorly; blade with about 25 deeply cut
teeth, rounded at the base. Terminal end of
shaft where it meets the blade with cusps.
Two to four heterogomph falcigers (Fig.
2C) below the spiniger, blades 5-10 pm
long, bidentate (first tooth much smaller
than second), with about five deeply cut
teeth; base rounded. Generally four falci-
gers per setiger from anterior to mid-body
setigers, then posteriorly diminishing to
two; blade length increases in mid-body
503
and then decreases again (Table 1). Outline
of single aciculum visible in some setigers;
details not clear. Ventral simple seta (base
width about 3 pm) in setigers 14—24; tip
strongly hooked and with short spines (Fig.
2D). Pygidium with two cirn, 0.13 mm
long and 0.03 mm long. Pharynx short, re-
tracted, extends forward from posterior por-
tion of setiger 2; single dorsally attached
tooth. Proventriculus 0.13 mm long and 0.1
mm wide, extends from setiger 2 to 5, with
about 13 muscle rows.
Description of San Salvador Island spec-
imens.—The following description is based
on 32 complete specimens and 13 attached
larvae, with 5—31 setigers, and ranging in
length from 0.34 to 2.7 mm and 0.09 to
0.22 mm in width; see Table 2 and Figs. 3—
5. Prostomium (Fig. 3A) about twice as
wide as long. Two pairs of dark red eyes
placed lateral to the antennae; anterior-
most, largest pair about half way along lat-
eral antennal length; posterior-most pair at
antennal bases. No obvious modification of
eyes in sexually mature specimens. Median
antenna 0.01 to 0.026 mm long and two
lateral antennae 0.01 to 0.015 mm long, all
short and originating slightly anterior to
prostomial-tentacular segment junction.
Tentacular segment about four times as
wide as long. Tentacular cirri 0.01 to 0.024
mm long (Figs. 3, 4). Digitiform dorsal
(0.02 to 0.03 mm long) and ventral (0.01 to
0.03 mm long) cirri on all setigers.
Setal structure and pattern similar for all
specimens examined. Dorsal simple seta
(2.4 to 3 wm base width) terminates bluntly
with about 15 short spines. Heterogomph
spiniger blade length 17 to 28 wm; blades
with approximately 30 teeth; spines on ter-
minal end of shaft. One to four hetero-
gomph falcigers (blade length 5 to 9 pm)
and spines on terminal end of shaft. One
ventral simple seta on posterior segments,
found in all setigers in attached larvae (6—
13 setigers) but are lost anteriorly in later
stages. Single, smooth aciculum (6 um bas-
al width) in each setiger, with rounded
slightly concave tip. Pygidium with two lat-
504 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
0.2 mM
Fig. 1. Dorsal view of Exogone breviantennata paratype (P-14590, ZMH) from El Salvador, 1.95 mm long,
0.2 mm wide, and with 24 setigers.
eral cirri (0.05 to 0.15 mm long), some-
times uneven in length, apparently due to
regeneration.
Pharynx (Figs. 3B, 4, 5) short, extends
to setigers 1, 2, or 3 depending on degree
of contraction; anterior rim with 10 soft pa-
pillae in addition to the large, curved, point-
ed, dorsally attached tooth, more easily
seen when pharynx is everted. Proventric-
ulus (0.07 to 0.2 mm long and 0.05 to 0.12
VOLUME 113, NUMBER 2 505
mS a eae
ia
a
eee
Fig. 2. Setae from Exogone breviantennata paratype (P-14590, ZMH) from El Salvador, 1.95 mm long, 0.2
mm wide, and with 24 setigers. A. Distal portion of dorsal simple seta, 3 4m basal width; B. Heterogomph
spiniger, 30 wm blade length; C. Heterogomph falciger, 10 zm blade length; D. Ventral simple seta, 3 4m basal
width.
506
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1.—Length in ym of spinigers and blades of compound setae from right side of Exogone breviantennata
paratype (P-14590, ZMH) from El Salvador, 1.95 mm long, 0.2 mm wide and with 24 setigers. utm = unable
to measure; avg = average; 0 = not present.
Setiger Heterogomph Heterogomph
number spiniger falciger#1
l 24 7.
2, 24 dL
3 30 Wf
4 30 10
») 30 10
6 32 utm
7-14 utm utm
15 24 7
16—20 utm utm
2A 20 5
22-24 utm utm
Averages 26415 TASS
mm wide) extends posteriorly from setigers
1—4, 2—4, 2-5, 2—6, 3-5 or 3—6 (Figs. 1, 5)
depending on degree of contraction during
fixation; about 15 rows of muscle cells.
Sexually mature females with internal
eggs or ventrally attached eggs or larvae
(Fig. 5). Six internal eggs, average diameter
0.08 mm in setigers 12—17 of a 25-setiger
female; five eggs 0.09 to 0.096 mm in di-
ameter, attached ventrally, one to two per
setiger on setigers 12, 14, and 15, of a 25-
setiger female; and six eggs 0.08 to 0.1 mm
in diameter, attached ventrally, two per se-
tiger on setigers 15, 19, and 22 of a 31-
setiger female. Five, 6-setiger larvae, (Fig.
5) average length and width 0.43 mm and
0.9 mm, one or two on setigers 12, 13, and
15 of a 21-setiger female; eight, 5-setiger
larvae, average length and width 0.26 mm
and 0.11 mm, one each on setigers 11-14
and 16—19 of a 27-setiger female. One 26-
setiger male found with sperm packed in
body cavity of setigers 18—25. No repro-
ductive modifications observed for either
sex. The only morphological differences
between juveniles and sexually mature
specimens are fewer setigers and more ven-
tral simple setae in the younger stages (Ta-
blew):
Discussion.—Two species of Exogone
that have a dorsal cirrus on the second se-
tiger and a short median antenna have been
Heterogomph Heterogomph Heterogomph
falciger#2 flaciger#3 falciger#4
Tl 6 5
Ti 6 5
a) 6 5
6 6 6
6 6 6
utm utm utm
utm utm 0
7 5 0
utm utm 0
5 0) 0)
utm utm 0)
6:35 5.80 55
reported from the Caribbean: E. brevianten-
nata and E. exmouthensis. A third species,
E. occidentalis, was not compared to E.
breviantennata when it was described from
the Galapagos Islands (Westheide 1974).
When Russell (1991) emended Westheide’s
(1974) description of E. occidentalis, based
on a study of the type material and on spec-
imens from a Belizean barrier reef, he de-
scribed most setae as bearing spination as
well as denticulation, thus bringing its de-
scription closer to E. breviantennata. Our
examination of the Belize specimens leads
us to agree with San Martin (1991) who
listed E. occidentalis as a junior synonym
of E. breviantennata.
We reexamined a paratype of E. brevian-
tennata and found that the setae showed the
spination and denticulation (also present in
Bahamian specimens), which were referred
to as indistinct in the type description. San
Martin (1991) figured the denticulation of
the blades of the compound setae and spi-
nation on the shafts on most setae for E.
breviantennata from Cuba.
Exogone exmouthensis was described
from Australia (Hartmann-Schroder 1980)
without comparison to E. breviantennata
and reported from Cuba by San Martin
(1991) without description. He placed E.
breviantennata and E. exmouthensis in sep-
arate subgenera based on details of the se-
VOLUME 113, NUMBER 2 507
Table 2.—Morphological measurements of specimens of Exogone breviantennata from Osprey Lake and
Reckley Hill Pond, San Salvador, Bahamas. Measurements in mm. Spec. = specimen; set. = setiger; provent.
= proventriculus; vent.SS = setiger number on which ventral simple setae begin; coll. = collected; loc. =
collection site; Loc. 1 = Osprey Lake; Loc. 2 = Reckley Hill Pond; utm = unable to measure. Figures for
specimens numbered 33-37 are averages from 5 larvae attached to specimen #4 and those for specimens 38—
45 are averages from 8 larvae attached to specimen #15.
Spec. Set. Body Body Provent. Provent. Vent. Date
# # length width length width SS coll. Loc. Comment
38-45 5 0.30 1.00 0.07 0.05 05/05/96 2 on spec. 15
33-37 6 0.43 0.09 0.07 0.05 ] 05/05/96 l on spec. 4
16 6 0.34 0.10 utm utm 1 06/19/97 Z
7 10 0.52 0.10 0.08 0.07 06/19/97 Z
18 10 0.70 0.10 0.08 0.05 ] 06/19/97 Z
19 12 0.63 0.12 utm utm 06/19/97 Z
20 12 0.71 0.12 0.10 0.06 8 06/19/97 Z
5 ie 0.85 0.17 0.12 0.09 4 05/05/96 l
21 13 0.70 0.11 Onz 0.07 1 06/19/97 pe
5 15 1.00 0.12 0.14 0.11 10 05/05/96 l
PPD 16 0.88 0.14 O13 0.07 10 06/19/97 2
1 16 1.20 On2 OS 0.1 9 05/05/96 ]
23 17 1.03 0.14 0.11 0.07 i 06/19/97 2
24 17 1.10 0.14 utm utm 10 06/19/97 2
2 A] 1.35 0.16 0.14 0.1 9 05/05/96 1
25 19 Lt 0.14 0.15 0.08 12 06/19/97 Ps with eggs
7 19 1.40 0.18 0.15 0.09 IZ 05/05/96
26 20 22 0.18 O17 0.10 utm 06/19/97 2
24 20 1.30 0.16 0.14 0.08 13 06/19/97 2
4 21 1.40 0.2 C5 Ont 22 05/05/96 l with larvae
12 2A 2.10 utm OA7 0.1 14 05/05/96 2
28 11 2.10 0.17 O11F 0.1 i) 06/19/97 2
9 22 13/5 utm 0.2 0.1 16 05/05/96 l
29 23 1.80 0.16 0.16 0.10 17 06/19/97 ou
13 pee) Dine utm 0.17 0.1 14 05/05/96 2
30 25 1.72 0.18 0.18 0.10 19 06/19/97 2
6 jib 130 utm 0.18 Or 18 05/05/96 l with eggs
51 25 2.00 0.16 utm utm 20 06/19/97 2
32 26 1.90 0.19 0.20 0.10 20 06/19/97 2 with sperm
8 pa | 1/95 0.20 O2 0.12 23 05/05/96 l
14 2d pIp 2.) 22 0.19 O12 23 05/05/96 2
15 pH Z.10 0.20 OF 0.1 22 05/05/96 2 with larvae
11 a1 ZS 0.20 0.18 0.1 18 05/05/96 ] with eggs
tae, e.g., head of the shaft of compound se-
tae with or without spination, the determi-
nation of which is dependent upon variables
unassociated with morphology of the spec-
imens themselves (see above). We have not
seen specimens of E. exmouthensis and so
are unable to comment on its relationship
to E. breviantennata.
Two other species, E. ovalis Hartmann-
Schroder, 1960, and E. breviantennata
ovalis Hartmann-Schroder, 1974, have been
made synonyms of E. breviantennata (San
Martin 1991).
Most species of the genus Exogone that
are described as having a dorsal cirrus on
setiger two also have a median antenna that
reaches at least to the tip of the prostomi-
um, if not longer. In this regard, E. brev-
iantennata is unique in that its median an-
tenna is much shorter, not even reaching to
the tip of the prostomium.
Because new species are sometimes
508 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
“VIZ
Fig. 3. Anterior end of Exogone breviantennata from Reckley Hill Pond, San Salvador, Bahamas. 1.72 mm
long, 0.18 mm wide, and with 25 setigers. A. Dorsal view of prostomium including first two setigers; B. Ventral
view including first setiger, showing anterior end of pharynx with tooth.
VOLUME 113, NUMBER 2
509
0.2 mm
Fig. 4. Exogone breviantennata from Osprey Lake, San Salvador, Bahamas. 1.4 mm long, 0.18 mm wide,
and with 19 setigers; lateral view of anterior end, showing tooth in extruded pharynx.
erected for minor setal differences, we care-
fully measured as many characters as pos-
sible for the paratype (from El Salvador)
and for specimens from Osprey Lake and
Reckley Hill Pond on San Salvador. We
found no significant differences from site to
site.
Habitat.—Type material collected in mud
around mangroves in El Salvador (Hart-
mann-Schroder 1959). In the Bahamas, E.
breviantennata was found amongst filaments
of the algae Batophora sp. and Vaucheria
sp. and on the prop roots of Rhizophora
mangle. Pascual et al. (1996) summarized its
habitat as intertidal to shallow depths, sand
and shell gravel, in sponges and amongst hy-
droids on R. mangle roots, Halimeda sp. in
beds of Thalassia testudinum, Lobophora
variegata and calcareous crusts, Vermetus
sp., photophilic algae and living inside the
cavities of six species of infralittoral demo-
sponges, where it constituted 0.6% of the to-
tal number of polychaetes and 9.7% of Ex-
ogone specimens extracted from the spong-
es. These habitats define it as part of an in-
terstitial and crevice fauna.
Distribution.—Equatorial Pacific: Gala-
pagos Islands (Westheide 1974), North East
Pacific: La Herradura, El Salvador (Hart-
mann-Schréder 1959); North Atlantic: Ba-
hamas (reported here), Belize (Russell
1991, as E. occidentalis), Canary and Ma-
deira Islands (Pascual et al. 1996), and
Cuba (San Martin 1991). San Martin (1991:
731) says “‘Probably circumtropical.”’
Biology.—The lack of solid material in
S10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
0.2mm
Fig. 5. Ventral view of Exogone breviantennata female with attached larvae from Osprey Lake, San Salvador,
Bahamas, 1.4 mm long, 0.2 mm wide, and with 21 setigers.
VOLUME 113, NUMBER 2
the digestive tract of the Bahamian speci-
mens suggests that they were fluid feeding,
most likely using their tooth to penetrate
algal filaments and then sucking out the
contents with the aid of the muscular pro-
ventriculus. See Figs. 3B and 4 for the tooth
in position for feeding. In live material, the
gut is often dark green along its entire
length, thus supporting this interpretation.
Pascual et al. (1996) reported that, in the
Canary and Madeira Islands, spicules of the
species of sponges from which specimens
were collected were found in their gut, sug-
gesting that the worms were actually eating
sponge tissue. There did not seem to be any
preterence for a particular species of
sponge. They speculated as to whether
polychaetes that are found with sponges
function at times as parasites, mutualists, or
commensals. Members of the subfamily
Exogoninae had been previously thought to
be selective deposit-feeders of mud or de-
tritus (Uebelacker 1984:30—4).
Reproduction.—San Salvadoran sexually
mature females were found with up to six
eggs or eight larvae (Fig. 5, Table 2) at-
tached to their ventral surface. Extrusion of
eggs from the body cavity and fertilization
were not observed. One male was found
with sperm. There was no evidence of re-
productive modifications of any kind. Hart-
mann-Schréder (1959) found gametes in,
and eggs attached ventrally on, setigers 11—
15 in the type material from El Salvador. In
Belize, Russell (1991) found a sexually ma-
ture female with oocytes in six setigers be-
ginning about setiger 16. Since attached lar-
vae from both San Salvador and Belize fe-
males (Russell 1991) had no more than six
setigers, they probably detach and become
benthic at this stage.
Previous Bahamian records of the genus
Exogone.—Worldwide, nearly 50 species
of the genus Exogone have been described
(Pascual et al. 1996). As of 1992, 19 of
these species that are considered valid have
been reported from the Caribbean and as-
sociated waters: E. arenosa Perkins, 1981;
E. atlantica Perkins, 1981; E. brevianten-
511
nata Hartmann-Schroder, 1959; E. cariben-
sis San Martin, 1991; E. dispar Webster,
1879; E. exmouthensis Hartmann-Schroder,
1980; E. gemmifera Pagenstecher, 1862; E.
longispinulata San Martin, 1991; E. lourei
Berkeley & Berkeley, 1938; E. naidina
@rsted, 1845: E. naidinoides Westheide,
1974; E. parhomoseta mediterranea San
Martin, 1984; E. pseudolourei San Martin,
1991; Exogone sp. A and sp. B, Uebelacker,
1984; E. rolani San Martin, 1991; sp. A,
San Martin, 1991; E. verugera Claparéde,
1868; and E. wolfi San Martin, 1991. See
Perkins & Savage (1975), Salazar-Vallejo
(1992) and Camp et al. (1998) for referenc-
es to Caribbean reports. Of these, two have
been reported from the Bahamas: Exogone
dispar (Webster) (based on Vittor & John-
son 1977 and our examination of USNM
51541, from Bimini Lagoon) and E. verug-
era (Claparéde) (in the collections of the
USNM, L. Ward, pers. comm., 29 Aug
1995). San Martin (1991) notes that the
original description of E. verugera is con-
fused, and therefore not reliable, and syn-
onymizes a series of Atlantic reports with
E. breviantennata. Seven of the species re-
ported from the Caribbean have a dorsal
cirrus on the second setiger: E. arenosa, E.
breviantennata, E. dispar, E. exmouthensis,
E. lourei, E. pseudolourei, E. rolani, and E.
wolfi. As any of the remaining six might be
found in Bahamian waters, they are includ-
ed in the key below.
Key to Caribbean Exogone species with
dorsal cirrus on second setiger
la. Median antenna does not reach tip of
prostomiumnisegas sok. cians i~.ce'u..
.. E. breviantennata Hartmann-Schroéder, 1959
1b. Median antenna, much longer than lat-
eral antennae, reaches to or exceeds tip
C50 RUSS COREA ORIN Goss no a pitt ns acer sw oc ee 2
2a. Tips of shafts of spinigers on setiger 2
with greatly enlarged triangular process
Pree hee eee oes eer ee eee i:
2b. Tips of shafts on setiger 2 not greatly
eilareed ax . seu 2). cs eee . +4
3a. Superior simple setae with well-defined
S12
spine on tips; proventriculus with up to
28 rows. of muscle cells. ssc, 4:6
E. arenosa Perkins, 1981
3b. Superior simple setae without spine;
proventriculus with at most 20 rows of
muscle cells: |. Were een pe eee ee
.... E. lourei Berkeley & Berkeley, 1938
4a. Dorsal simple setae of median and pos-
terior setigers enlarged and modified,
strongly bidentate; proventriculus short,
with ca 15 muscle:cells rows ......
E. pseudolourei San Martin, 1991
4b. Dorsal simple setae of median and pos-
terior setigers not enlarged
5a. Median antenna club shaped ........
ie ste Se: E. dispar (Webster, 1879)
7 2e © © © © © © © ew ee
bins eh er eye
Sb. Median antenna elongate 2. ..5 s..o5< . 6
6a. Dorsal simple setae similar throughout
Ody chases. E. wolfi San Martin, 1991
6b. Dorsal simple setae increasing in thick-
ness posteriorly and changing in shape
i. 4 Te E. rolani San Martin, 1991
Acknowledgments
We were alerted to the interesting poly-
chaete fauna of the land-locked marine
lakes of San Salvador, The Bahamas, by
William Lindsay (Elmira College, New
York), who, along with the staff of the Ba-
hamian Field Station, have made this work
possible. We thank them and Fitchburg
State College for its continued support. We
especially appreciate the constructive com-
ments on the manuscript by Drs. Nathan
Riser and David Russell.
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):514—525. 2000.
Podarke aberrans Webster & Benedict, 1887 - resolution, with
descriptions of two new species in the genus Microphthalmus
(Annelida: Polychaeta)
Nathan W. Riser
Marine Science Center, Northeastern University, Nahant, Massachusetts 01908, U.S.A.
Abstract.—Reference is made to individuals with ‘“‘fan-shaped”’ caudal ap-
pendages from two different regions in the Gulf of Maine and to two different
life-history stages in the original description of Microphthalmus aberrans
(=Podarke aberrans Webster & Benedict, 1887). Intertidal meiofaunal inves-
tigations in the region over the past forty years have yielded three species that
correspond to parts of the description. Two fragments of the ‘“‘adult form”
labeled Type, from Eastport, Maine, upon which most of the type description
is based, exist as one of the syntype slides deposited by Webster & Benedict
in the National Museum of Natural History. The species has been encountered
occasionally in clean coarse sand beaches along the coast of Maine, and in the
subtidal of Nahant Bay, Massachusetts. A second species with a fimbriate anal
lamella, M. aggregatus n.sp., occurs in the intertidal of Cape Cod Bay, Mas-
sachusetts. The “‘half grown specimens”’, M. pettiboneae n.sp., to which the
epithet aberrans has been incorrectly applied, is also present among the syntype
slides, and has been collected in intertidal samples from New Brunswick, Can-
ada to the south shore of Cape Cod, Massachusetts.
Webster & Benedict (1887) assumed that
badly damaged specimens from Province-
town, Massachusetts collected in 1879, but
not retained, were the same species as the
one being described as Podarke aberrans
from the Eastport area in Maine, because of
the “peculiar fan-shaped’? anal lamella.
One “‘adult’’ specimen was encountered by
the authors at Eastport, and the description
of Microphthalmus abberans is based upon
that individual with additional remarks on
the morphology of “‘half-grown speci-
mens.” The figures accompanying the de-
scription are of both the ‘adult’? and ju-
venile form, and the type material of Mi-
crophthalmus aberrans (Webster & Bene-
dict, 1887) (=Podarke aberrans W. & B.)
deposited in the National Museum of Nat-
ural History (USNM), is a mixture of two
species. Southern (1914) reported that he
had examined the “original types”? depos-
ited in the USNM and at Union College and
that they did not agree with the type de-
scription. He transferred the species from
Podarke to Microphthalmus, accepting the
specific epithet aberrans for the material
available to him and furnished a figure of
the notopodial setae to support his conten-
tion; thus, we must assume that the slide of
the specimen upon which Webster & Ben-
edict based their description was not en-
countered. Eliason (1920) accepted South-
ern’s statements about the type material
while describing material from the Ore-
sund, which he ascribed to the Webster &
Benedict species, thus establishing a pre-
cedent of applying the epithet aberrans to
a species that does not agree with the type
description. The failure of subsequent au-
thors to examine the type material has re-
sulted in a perpetuation of this error.
A few individuals fitting the description
of Microphthalmus abberans have occurred
in intertidal meiofaunal collections I have
VOLUME 113, NUMBER 2
made along the coast of Maine over the past
forty years and in subtidal substrate from
Nahant Bay, Massachusetts. Numerous
specimens of a second species with a fim-
briate anal lamella have routinely been en-
countered in the intertidal dunes on the sand
flat at Ellisville, Massachusetts, as well as
on the beach at Manomet, Massachusetts
across Cape Cod Bay from Provincetown.
The initial description of Podarke aberrans
refers to three different species belonging
to the genus Microphthalmus, two of
which, including the species that has born
that name since Southern’s publication,
must be described as new.
Materials and Methods
Specimens were obtained from substrate
collected in 18 fl. oz. plastic bags for mei-
ofaunal studies. Each sample was washed
with fresh sea water in the laboratory and
decanted onto 153 wm screens from which
animals were removed for sorting. The sed-
iment was then extracted with 7.5% MgCl,
and decanted onto the screens from which
the animals were washed into fresh sea wa-
ter. Specimens were anaesthetized with
7.5% MgCl, prior to fixation in Hollande’s
cupri-picri-formal-acetic. Whole mounts
were Stained with Ranvier’s picro-carmine
or Mayer’s alcoholic HCl carmine; some
were counterstained with alcoholic indigo-
carmine. All measurements were obtained
from living specimens.
Material labeled Microphthalmus aber-
rans in the collections of the Atlantic Ref-
erence Centre (ARC), St. Andrews, New
Brunswick, and the National Museum of
Natural History were obtained on loan. The
syntypes (USNM 447) of Podarke aberrans
from Eastport, Maine deposited by Webster
& Benedict consist of five slides and two
specimens in alcohol.
Microphthalmus aberrans (Webster &
Benedict, 1887)
Figs. 1-6
Podarke aberrans Webster & Benedict,
1887:713-—715, p.p., “‘adult form’’, figs. 14,
515
15, 18. Microphthalmus aberrans.—Petti-
bone, 1963:104, p.p.
Diagnosis.—Sexual individuals to 9 mm
in length with 30—45 setigers. Ocelli ab-
sent. Dorsal cirri more than four times as
long as neuropodial lobe. Anal cirri slightly
longer than dorsal cirri. Notopodia with up
to 15 pointed simple and one pectinate seta
and a sturdy aciculum. Presetal neuropodial
ligule with thin aciculum associated with
one or two bidentate simple setae (Fig. 3).
Neuropodial setal bundle with thick acicu-
lum, one apically bidentate simple seta and
about 15 finely denticulated apically biden-
tate falcigers. Falciger blades range from 6—
32 wm in length. Testes in setigers 9
through 16, ovaries in following setigers.
Penis anterior to parapodia of setiger 3 on
both sides, with tubular stylus 25 wm long,
muscle bulb large with large posterior pa-
pilla. Anal lamella with about 30 short,
blunt, fimbriae.
Lectotype.—USNM (447), slide 502, la-
beled “‘Podarke aberrans adult fragments”
(two fragments; one of three setigers from
the female region of the body, and one of
four setigers and anal lamella).
Material examined.—Data and _ photo-
graphs obtained from living specimens col-
lected in the intertidal at Liberty Point,
Robbinston, Maine; Crowe Neck, Cobs-
cook Bay, Maine; Griffith’s Head, George-
town, Maine, and 16 m depth off Egg Rock,
Nahant Bay, Massachusetts, in addition to
the lectotype slide, have made it possible to
complete the description of the species. One
adult (USNM 186552) collected on 27 Mar
1980 at Liberty Point and one immature
specimen (USNM 186533) encountered at
Griffith’s Head on 20 Apr 1999, have been
deposited, in alcohol, in the USNM.
Morphological notes.—The acicula taper
toward the slightly expanded apex; that of
the presetal neuropodial lobe is hooded.
The diameter of the presetal neuropodial
aciculum is almost one-half that of the other
two, <3 vs. 6 wm, however it runs at an
angle, and its base varies in position from
the same level to 4 wm distal to that of the
516
neuropodial aciculum. The acicula are all
approximately the same length, 76 wm. The
pectinate notopodial seta ranges from 91-—
99 wm in length with the blade occupying
almost one-half of the length. The blade is
finely denticulated almost to the tip (Fig. 6).
The difference in construction of the blade
and shaft allows for great flexibility at the
juncture so that at times the seta takes on
the appearance of being jointed; the twisted
appearance figured for the pectinate seta of
M. bermudensis Westheide, 1973, (Abb. 4,
fig. D) has also been encountered. The pec-
tinate setae of species with several simple
notopodial setae can rarely be recognized in
the clump (Fig. 2) but can usually be dis-
tinguished in some of the parapodia of liv-
ing specimens under coverslip pressure.
The simple notopodial setae vary in ap-
pearance, depending upon the angle at
which they are viewed and if they are re-
tracted into the body or are projected. The
majority of the simple notosetae of pre-
served specimens are strongly hooked api-
cally. The striated fibrillae forming the cor-
tex spiral (180° ?) but can not be traced to
the apex. This spiraling may account for the
flexibility which results in the hooked con-
figuration.
There appear to be eight digitiform pa-
pillae on the pharynx cap. The pharynx ex-
tends posteriorly to setiger 5 which is oc-
cupied by the ventriculus. The epithelium
of the pharynx is thick and when the organ
is inflated, develops longitudinal ridges
(Fig. 1). There is a well-developed valve
and sphincter separating the pharynx and
ventriculus. The latter is globular and com-
prised of radial muscle and glandular cells.
Retractor muscles extend from the base of
the pharyngeal cap to the body wall at the
juncture of the third tentacular segment and
first setiger at which point protractors orig-
inate passing to insert at the juncture of the
pharynx and ventriculus. Strongly devel-
oped retractor muscles extend from the ven-
triculus to the longitudinal muscles of se-
tiger 8.
One of the 18 sexually mature specimens
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
had an ovary on the left side, and testis on
the right of setiger 16.
Remarks.—tThe figure of the anal lamella
in the description by Webster & Benedict
has little relationship to the specimen on the
lectotype slide, except that the midregion is
different from the rest of the margin. The
lamella of living animals (Fig. 5) is domed,
and if not fully expanded projects over the
fimbriae so that they may appear to be ab-
sent medially for a variable extent of the
margin (Fig. 4). A fold in the middle of the
dorsal surface of the anal lamella on the
lectotype may have been interpreted by
Webster & Benedict as corresponding to the
anal lamella of the “half grown’’ speci-
mens.
Pettibone (1963) described the general
appearance of the notopodial simple setae
on the adult slide of the type specimen of
M. abberans but did not report observations
on notopodial setation of other specimens
which she identified as that species (some
of which were mature). The simple noto-
setae of preserved specimens of M. fragilis
Bobretzky, 1870 are thin and sharply point-
ed; some are slightly bent toward the apex;
those of M. urofimbriatus Alikunhi, 1948
were reported to be straight. [Hartmann-
Schr6der (1960) stated that the setae of M.
c.f. urofimbritta (sic) from the Red Sea
were too delicate to allow for characteriza-
tion. The clusters of notopodial simple setae
present in M. aberrans (Fig. 1), M. fragilis,
and M. urofimbriatus are so apparent that it
is more likely that her worms belong to the
M. similis group of species.] The apex of
the pectinate seta of M. fragilis is long and
slender beyond the denticulated portion of
the blade.
The species has not been routinely en-
countered nor in significant numbers. The
single specimen obtained from the subtidal
of Nahant Bay was associated with Ophry-
otrocha cf. gracilis Huth 1934, Parougia
caeca (Webster & Benedict, 1884) and an
eyeless species of Protodorvillea, all sub-
tidal species in Nahant Bay but routinely
VOLUME 113, NUMBER 2 517
j
Bee ae ie tr had
-# yO te 3
Pe Lea eee oes
a
Sa. tat
ate ey
Figs 1-10. 1—6; Microphthalmus aberrans, living specimens. |. Optical section of foregut region. 2. Noto-
podial setal bundle in situ. 3. Paired simple neuropodial seta in situ. 4. Anal lamella, anal cirri detached, dorsal
view. 5. Anal lamella second specimen, anal cirri detached, ventral view. 6. Blade of pectinate seta, dissociated
preparation. 7-9; M. aggregatus, 7. Optical section of foregut region. Living specimen. 8. Anal lamella and anal
cirri, living specimen. 9. Penis stylet and papilla, living specimen. 10; M. pettiboneae. 19 «wm falciger blades,
slightly oblique. Fixed specimen.
518
present in the intertidal of the coast of
Maine.
Microphthalmus aggregatus, new species
Figs. 7-9, 11-15
?Podarke aberrans Webster & Benedict,
1887:713 p.p., ““same form at Province-
town, Massachusetts.”’
?Microphthalmus c.f. similis Westheide &
Rieger, 1978; Westheide, 1979.
Diagnosis.—Transparent except for
opaque white gonads. Individuals with go-
nads 3.2—7.1 mm long; with 31—43 setigers.
Paired and median prostomial antennae ap-
proximately same length; palps extend
about same distance beyond prostomium as
paired tentacles. Blackish brown ocellus
about mid-way between bases of paired
prostomial antennae and Ist tentacular cir-
rus on each side. Dorsal cirri about four
times length of neuropodium. Notopodium
of first setiger with aciculum, following se-
tigers with notopodial aciculum and pecti-
nate seta. Neuropodium with bidentate fal-
cigers and one or more simple setae. Phar-
ynx bipartite; ventriculus oval to round, oc-
cupying 3rd setiger. Testes bilobed, in
setigers 9 through 16; ovaries in subsequent
setigers to penultimate. Penis digitiform
with eight stylets and dorsal sensory papilla
(Fig. 9); anterior to parapodia of third se-
tiger on each side. Anal lamella with digi-
tiform fimbriae.
Holotype.-—USNM_ 186534. Massachu-
setts; Ellisville, intertidal coarse sand.
Paratypes.—USNM_ 186535, 12 speci-
mens in alcohol. Peabody Museum of Nat-
ural History, Yale University, New Haven,
Connecticut: YPM 24000, YPM 24001,
stained whole mounts.
Etymology.—Latin aggregatus, clus-
tered; referring to the tendency to occur in
large concentrations, and to aggregate when
isolated in vitro.
Material examined.—More than two
hundred specimens from Ellisville, Massa-
chusetts have been collected. Morphologi-
cal data were obtained from 84. Additional
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
data were recorded from specimens collect-
ed on Manomet Beach, Massachusetts
(about 1 km North of Ellisville).
Morphological notes.—The body tapers
from the anterior end to the female region,
but the interparapodial width is about con-
stant. Bands of refringent granules in the
dorsal epidermis are colorless and do not
produce opacity. Pigment is restricted to
black granules in the epithelial cells of the
rectum. The setigers containing ovaries are
inflated and have a larger diameter than the
anterior setigers. The prostomial antennae
and palps range from 0.13—0.15 mm in
length; the median antenna is thinner and is
occasionally forked. The palpophore is
small, accounting for very little of the ex-
tent of the palp. The dorsal tentacular cirri
are longer than the ventral, viz., lst 0.20—
0.22:0.13—0.16, 2nd 0.27:0.14-0.16, 3rd
0.27—0.30:0.05—0.07 mm. Dorsal cirri of
parapodia 0.24—0.27 mm long, except setig-
er 1 which are shorter, rarely exceeding
0.17 mm. Frequently, apparently as the re-
sult of regeneration, individual tentacular
and dorsal cirri are shorter than these mea-
surements. The pleura of all setigers expand
laterally and the cirrophores are elongate,
capped distally with a lobe above the base
of the cirrus (Fig. 11). This lobe contains
numerous mucous cells. The pair of anal
cirri to either side of the anus attain lengths
up to 0.4 mm. (Fig. 8). The fully expanded
anal lamella is about 0.25 mm broad with
as many as 42 digitate fimbriae 0.03 mm
long. Shorter fimbriae as well as those that
are forked probably indicate injury.
Ciliation on the prostomium is restricted
to the nuchal organs (Fig. 11), each of
which bears a small papilla on the anterior
margin. A band of cilia extends across the
dorsum between the bases of the second
tentacular cirri but is interrupted medially
on the subsequent four segments. Complete
ciliary bands extend from the median edge
of the lobe at the apex of the cirrophore
(Fig. 11) across the dorsum of succeeding
setigers. There is a lateral cluster of cilia
anterior to each parapodium, and a lateral
VOLUME 113, NUMBER 2 319
*
Figs. 11-14. Microphthalmus aggregatus, SEM. 11. Dorsal view, anterior end. 12. Ventral view, anterior
end. 13. Anal lamella, ventral view. 14. Medium and short falciger blades.
520
patch posteriad. Ventrally, a band of cilia
extends along the posterior edge of the
mouth (Fig. 12), constituting the only con-
sistent ciliation on the ventral surface.
Cirrophores of the dorsal cirri are almost
as long as the neuropodia. Notopodial lobes
are not present. Notopodial setation consists
of one or two pointed acicula 45-57 pm
long and one or two pectinate setae 24—33
wm long of which the comb occupies one-
fourth to one-third of the length. The pre-
setal lobes of the neuropodia are conical,
about 22 ym long and are supported by a
thin pointed aciculum 57 wm long accom-
panied by a bidentate simple seta which
moves independently, or sometimes in con-
junction with the other neuropodial setae.
A pointed simple or bidentate seta, or both,
may be present in the neurosetal bundle,
which consists of as many as nine apically
bidentate falcigers (Fig. 14), most of which
are 17—20 pm long and accompanied by
one or more with short blades 7-8 wm long
and usually one or more with blades in ex-
cess of 36 wm long. Ventral cirri of the
parapodia are thin and filiform ranging be-
tween 35 and 42 um in length.
The soft portion (cap) of the pharynx oc-
cupies the first tentacular segment and bears
eight simple digitiform papillae tightly
packed together around a large and expan-
sive surface. Retractor muscles pass directly
from the base of the cap to the body wall
of the third tentacular segment from
whence, protractors extend to the base of
the pharynx. The second region occupies
the second tentacular segment and is mus-
cular (Fig. 7) with an epithelium containing
granular products. The third region extends
from the third tentacular segment to the
third setiger. The ventriculus is firmly at-
tached to the adjacent wall of the body.
Remarks.—In April and May, the ovaries
are packed with yolky oocytes 35 wm in
diameter, and the testes are primarily in ear-
ly stages of spermiogenesis with few sper-
matids undergoing flagellar development.
By August, the testes are packed with fully
developed sperm; ovarian segments contain
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
sperm in seminal receptacles; and the num-
ber of oocytes is reduced. Ovaries contain
two or three degenerating oocytes in No-
vember and December, but the testes are
packed with sperm. The animals have lived
in the sorting bowls for up to eight months
with biweekly water changes, but egg lay-
ing has not occurred in the absence of a
food source, and the gonads have been re-
sorbed. Efforts to investigate posterior re-
generation, as the result of the discovery of
a specimen with a bifurcated posterior end
(Fig. 15), in 1977, have been futile for the
same reason.
The major difference between M. aggre-
gatus and M. similis Bobretzky, 1870, in
addition to size and number of segments, is
the distribution of the gonads. Ovaries of
M. similis do not occur anterior to setiger
20. About 10% of the M. aggregatus indi-
viduals (7 of 92) had the ovary on the right
side of setiger 16 and testis on the left, and
the first testis on the right in setiger 8 in-
stead of 9. One specimen was encountered
with a mixed gonad in the right antimere of
setiger 17 and ovary on the left. Some spec-
imens of M. aggregatus have lacked pig-
mented ocelli, while others have lacked
them on one side, or have had two (one
dorsal and one ventral) on one or both
sides. Variation, as well as absence of ocelli
in M. fragilis, was noted as an intraspecific
variation by Bobretzky (1870).
The divided pharynx of M. fragilis fig-
ured by La Greca (1950:fig. 1) was not ap-
parent in any of the three specimens from
the Black Sea which I examined, but the
pharynx of M. similis from there was all
divided as in M. aggregatus.
Microphthalmus pettiboneae, new species
Fig. 10
Podarke aberrans Webster & Benedict,
1887: 713-715, p.p. ‘“‘juvenile form’’,
fies: 16517, 19; 20:
Microphthalmus aberrans Southern, 1914:
46, fig. 7._Pettibone, 1963:104, p.p., figs.
27a—b.—Clausen, 1986:184—186, figs.
VOLUME 113, NUMBER 2
pAC5
16
Figs. 15,16. 15. Microphthalmus aggregatus, with posterior bud, living specimen. Scale equals 0.09 mm. 16.
Anal lamella of living Microphthalmus nahantensis Westheide & Rieger, 1987. Scale equals 0.045 mm.
14, 19 (for additional synonymy see
Hartmann-Schroder 1971).
Diagnosis.—Variable intensity of brown
pigmentation; mature individuals less than
10 mm in length with 30-35 setigers; pair
of ocelli present; dorsal cirri only slightly
longer than neuropodial lobes; unpaired
sucker-like penis; single pectinate and sim-
ple seta with an aciculum in notopodium;
testes in setigers 6 through 9; bilobed (me-
dially indented) anal lamella.
Syntypes.—USNM (447), slides 137,
138, 501, and two in alcohol deposited by
Webster & Benedict.
Additional material deposited.—USNM
(186536) from Beacon Point, Blacks Har-
bour, New Brunswick, Canada: ARC
9953141; 995314, from Pagan Point, St.
Andrews, New Brunswick, Canada.
Etymology.—The species is named for
Dr. Marian Pettibone, who in 1953, was re-
sponsible for my initial efforts to resolve
the problem.
Material examined.—Observations have
been recorded from numerous living indi-
viduals collected in intertidal coarse sand
beaches between Beacon Point, Blacks Har-
bour, New Brunswick, Canada and East
Point, Nahant, Massachusetts, USA. Spec-
imens identified by Pettibone as M. aber-
rans (USNM 28413) from Southport Island,
Maine; (USNM 32498) from Rye Beach,
New Hampshire, and from Halifax, Nova
Scotia, Canada (USNM 49180) were ex-
amined and found to be M. pettiboneae, as
were specimens from the subtidal (ARC
9661230) of Lime Kiln Bay, Letang Estu-
ary, New Brunswick, and (ARC 9953146)
“off’? Navy Island, St. Andrews, New
Brunswick in the Atlantic Reference Centre
collection.
Morphological notes.—The simple no-
topodial seta tapers to a point, and is strong-
ly developed, approximately 51 wm long
and slightly more than 2 wm maximum di-
ameter. The cutting edge of the pectinate
seta is reinforced by 8—12 strongly devel-
oped denticles, beyond which the blade is
thin and serrations are not visible. Blades
of the neuropodial falcigers are bidentate
with serrated margins; one or two with fine
denticulations are long and spiniger-like,
522 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
measuring 75-85 wm in length; three to
five, with strong denticulations, are about
40 wm; two with strong denticulation are
small, about 12 wm long; while some (Fig.
10) approximate 20 wm in length. One or
two simple neuropodial setae are present in
the terminal 2—4 setigers.
Remarks.—Westheide (1967) noted that
the juveniles included in the type descrip-
tion of M. abberans constituted a separate
species characterized by the presence of
eyes, short cirri, and a simple rounded anal
lamella, features for which no ontogenetic
evidence supported the possibility of
change to the “‘adult’’” morphology.
Denticulation of the blades of the pecti-
nate setae of M. pettiboneae is difficult to
show accurately in line drawings, and while
the SEM micrograph of this seta in Clausen
(1986) from a North Sea specimen, is
slightly rotated, denticulation is similar to
that of the New England forms. The strong-
ly developed simple notopodial seta fre-
quently projects from the body of preserved
specimens and is the primary morpholog!-
cal feature by which immature M. pettibo-
neae can be distinguished from M. sczel-
kowii (Table 1). Mature specimens of the
two species are easily distinguished by o0o-
cyte size and number (Table 1).
M. scezelkowii
<2 neuropodial cirri
hemisphere
paired
>200 wm
<2
present
9/10
M. pettiboneae
<2 neuropodial cirri
hemisphere
single
<100 wm
many
present
9/10
M. nahantensis
>3%X neuropodial cirri
paired
>200 wm
<=13
spatulate
l
9/10
absent
M. aggregatus
>3 neuropodial cirri
present
fimbriate
paired
<100 pm
many
16/17
Discussion
The genus Microphthalmus is hermaph-
roditic and its members are typical met-
ofaunal animals attaching to hard particles
with a caudal adhesive organ and coiling
when disturbed. Intact specimens are infre-
quently present in routine screenings of
substrate, and pretreatment to release at-
tached animals is essential in order to ob-
tain complete specimens or population den-
sities.
An investigation similar to that of West-
heide & Rieger (1987), which clarified the
systematics of the M. listensis group of spe-
cies, may demonstrate that the species listed
in the synonymy of M. aberrans by Hart-
mann-Schréder (1971) are M. pettiboneae
M. abberans
>3X neuropodial cirri
absent
fimbriate
tor ls
paired
<100 wm
Table 1.—Morphological features of New England species of Microphthalmus.
many
16/17
Notopodial simple setae
Dorsal cirri
Penes
Oocyte diameter
Anal lamella
Oocyte number
M/f juncture
Ocelli
VOLUME 113, NUMBER 2
or that speciation has occurred and they
constitute different species.
Very young individuals of M._ similis
were described by Westheide (1967) from
the intertidal at Sylt and Hartmann-Schréd-
er & Stripp (1968) recorded sexually im-
mature individuals up to 18 mm long and
consisting of more than 60 segments, char-
acteristics which agree with those of M.
similis, from a depth of 21 m in the North
Sea. Gonad distribution and penis charac-
teristics have not been reported for the
North Sea populations. The specimens list-
ed as M. cf. similis by Westheide & Rieger
(1978) from North Carolina, U.S.A. were
small and immature. The distribution of go-
nads in the two sexually mature specimens
from the same region was not recorded by
Westheide (1979). It is possible that they
represent a southern distribution of M. ag-
gregatus or constitute a separate species.
Westheide (1977) noted that reproductive
organs were significant but could not be in-
cluded in a phylogenetic analysis of Mi-
crophthalmus because of the absence of
data for many species. However, the male-
female (m/f) juncture of the five New Eng-
land species (Table 1) supports the possible
adelphotaxa indicated in his cladogram.
The occurrence of ova in one antimere and
sperm in the other of setigers at the m/f
juncture of M. aggregatus has also been re-
corded by Bobretzky (1880) for M. fragilis
and M. similis and Clausen (1896) for M.
cf. pettiboneae.
The prostomium of Microphthalmus spe-
cies is malleable, influencing its shape and
the location of the palps and antennae. Re-
traction of the prostomium into the peris-
tomium may draw the median antenna back
against or into the peristomial fold. The me-
dian antenna is thin, rarely visible from
ventral view unless bent to one side, and is
often undetectable on preserved specimens.
Ciliation has been described and figured
for members of the Jistensis-group by
Westheide & Rieger (1987), but other than
for the present description of M. aggregatus
523
has not been recorded for other species in
the genus.
A pectinate notopodial seta is character-
istic of members of the genus but is absent
in M. hartmanae Westheide, 1977 and M.
simplicichaetosus Westheide & Purschke,
1992, species with fimbriate anal lamellae,
dorsal cirri slightly more than twice as long
as the neuropodium and in which strongly
serrated simple setae are present in the neu-
ropodium (Westheide 1977, Westheide &
Purschke 1992). The pectinate notopodial
setae may be involved in stimulating the
glands on the cirrophore or in spreading se-
cretions from those glands. The role of no-
topodial setae in members of the genus with
more than one or two simple setae in ad-
dition to the comb seta is difficult to assess.
The presence of the male copulatory or-
gans in the third setiger is a generic char-
acter; however, the morphology and num-
ber of penes varies between species. Trans-
mission electron microscopy (TEM) by
Westheide (1979) demonstrated that the pe-
nis stylets of M. cf. similis from North Car-
olina are separate entities and not ridges in
the wall of the penis.
The presence of an aciculum in the pre-
setal neuropodial lobe appears to be char-
acteristic of the Microphthalminae sensu
Hartmann-Schroder (1971) while absent
from that lobe in the Hesionidae s.str.
Westheide (1977), with ‘‘hesitation’’,
postulated the fimbriate anal lamella as
“‘ancestral’’. Dorsal cirri of the parapodia
more than twice as long as the neuropodium
appears to be characteristic of species in
which the anal lamella is a large free fan as
in the fimbriate species and listensis-group
(Fig. 16). [The figure and description of the
anal lamella of M. c.f. urofimbritta (sic) by
Hartmann-Schréder (1960) is of a pre-
served specimen with the lamella flexed so
that the anal cirri and anus are terminal, and
the fimbriate posterior margin has folded
back beneath the terminal setigers. Artistic
license is also apparent in the figure of the
anal lamella of M. similis in La Greca
524
(1950) and M. abberans in Webster & Ben-
edict (1887).]
Gland openings on the anal lamella of
fimbriate species are restricted to the ventral
surface of the fimbrae (Fig. 13). The ar-
rangement of these glands has not been re-
corded for either the Jistensis-group nor for
the species with a simple hemispherical
(sometimes indented) lamella, and thus is
not available for assessment at present.
The ventriculus is attached by muscle fi-
bers to the body wall, and is restrained pos-
teriorly by the septum at the anterior end of
the intestine. Telescoping of segments alters
the topography as does eversion of the
pharynx. Eversion in species with a bipar-
tite pharynx (including the related genus
Hesionides) does not significantly alter ven-
tricular location; which is in setiger 2 in the
listensis-group, 3 in a number of species, 5
in aberrans, [the pharynx was reported by
Westheide & Purschke (1992) to extend to
setiger 4 in M. simplicichaetosus, which
would place the ventriculus in 5], 12 in M.
hamosus Westheide, 1982. The location of
the ventriculus, or origin of the intestine,
have not been routinely recorded in descrip-
tions of species.
Acknowledgments
The constant advice and encouragement
of Drs. Pettibone and Westheide throughout
the years would have been in vain if I had
not finally checked the type material of M.
aberrans deposited in the USNM (courtesy
of William Moser). The collection of Mi-
crophthalmus species present at the Atlantic
Reference Centre was obtained through the
courtesy of Dr. G. Pohle. Translations of the
two papers in Russian by Bobretsky were
furnished by Dr. Pettibone. Alcoholic spec-
imens of M. fragilis and M. similis from the
Black Sea were generously furnished by Dr.
T. Marinov in 1979 for comparative pur-
poses. The manuscript was greatly en-
hanced by the conscientious reviews of Drs.
Brigitte Hilbig and Stephen Gardiner.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Literature Cited
Alikunhi, K. H. 1948. On a new hermaphrodite species
of Microphthalmus (Polychaeta-Hesionidae)
from the sandy beach, Madras.—Journal of the
Royal Asiatic Society of Bengal. Science 14:
17-25.
Bobretsky, N. 1870. On the fauna of the Black Sea.
(In Russian).—Zapiski Kievskago Obshchestve
Estestvoispytatelei 1:188—274.
. 1880. Kutschenijn ob organach rasmnoschen-
ija u annelid.—Zapiski Kievskago Obshchestve
Estestvoispytatelei 6:61—86.
Clausen, C. 1986. Microphthalmus ephippiophorus
sp.n. (Polychaeta:Hesionidae) and two other
Microphthalmus species from the Bergen Area,
Western Norway.—Sarsia 71:177—191.
Eliason, A. 1920. Biologisch-faunistische Untersu-
chungen aus dem Oresund. V. Polychaeta.—
Lunds Universitets Arsskrift, Avd. 2, 16(6):1—
103.
Hartmann-Schroéder, G. 1960. Polychaeten aus dem
Roten Meer.—Kieler Meeresforschungen 16:
69-125.
1971. Annelida, Borstenwiirmer, Polychae-
ta.—Tierwelt Deutschlands 58:1—594.
, & K. Stripp 1968. Beitrage zur Polychaeten-
fauna der Deutschen Bucht.—Veroffentlichun-
gen des Instituts fiir Meeresforschung in Bre-
merhaven 11:1—24.
La Greca, M. 1950. Sulla presenza nel Mediterraneo
di Lycastoides pontica (Bobr.), Microphthalmus
fragilis (Bobr.) e Microphthalmus similis
(Bobr.).—Annuario dell’Istituto e Museo di
Zoologia della UnversitA di Napoli 2:1—16.
Pettibone, M. H. 1963. Marine polychaete worms of
the New England region. 1. Aphroditidae
through Trochochaetidae.—Bulletin of the Unit-
ed States National Museum 227:1—356.
Southern, R. 1914. Archiannelida and Polychaeta, in
Clare Island survey, pt. 47.—Proceedings of the
Irish Academy Dublin 31:1—160.
Webster, H. E., & J. E. Benedict. 1887. The Annelida
Chaetopoda from Eastport, Maine.—Report of
the United States Commissioner of Fisheries
1885. Pp. 707-755.
Westheide, W. 1967. Monographie der Gattungen He-
sionides Friedrich und Microphthalmus Me-
cznikow (Polychaeta, Hesionidae). Ein Beitrag
zur Organisation und Biologie psammobionter
Polychaeten.—Zeitschrift fiir Morphologie der
Tiere 61:1—159.
. 1973. Zwei neue interstitielle Microphthal-
mus-Arten (Polychaeta) von den Bermudas.—
Mikrofauna Meeresboden 14:1—16.
. 1977. Phylogenetic systematics of the genus
Microphthalmus (Hesionidae) together with a
description of M. hartmanae nov. sp. Pp. 103—
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113 in D. J. Reish & K. Fauchald, eds., Essays
on polychaetous annelids in memory of Dr.
Olga Hartmann.—Allan Hancock Foundation
Special Publication.
. 1979. Ultrastruktur der Genitalorgane inter-
stitieller Polychaeten II. Mannliche Kopulation-
sorgane mit intrazellularen Stilettstaben in einer
Microphthalmus-Art.—Zoologica Scripta 8:
111-118.
, & G. Purschke 1992. Microphthalmus simpli-
cichaetosus (Annelida:Polychaeta), a new he-
sionid from the Northwestern American Pacific
a20
Coast with exclusively simple chaetae.—Pro-
ceedings of the Biological Society of Washing-
ton 105:132-135.
, & R. M. Rieger 1978. Cuticle ultrastructure
of hesionid polychaetes (Annelida).—Zoomor-
phologie 91:1—18.
, & . 1987. Systematics of the amphi-
atlantic Microphthalmus-listensis-species-group
(Polychaeta:Hesionidae): Facts and concepts for
reconstruction of phylogeny and speciation.—
Zeitschrift fiir zoologisches Systematik und Ev-
olutionsforschung 25:12—39.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):526—531. 2000.
Parapionosyllis cabezali, a new species of Exogoninae
(Polychaeta: Syllidae) from Spain
Julio Parapar, Guillermo San Martin, and Juan Moreira
(JP) Departamento de Bioloxia Animal, Bioloxia Vexetal e Ecoloxia, Facultade de Ciencias,
Universidade da Corufia, Campus da Zapateira, E-15071 A Coruna, Spain;
(GSM) Laboratorio de Biologia Marina e Invertebrados. Departamento de Biologia, Unidad de
Zoologia, Facultad de Ciencias, Universidad Aut6noma de Madrid, Canto Blanco, E-28049
Madrid, Spain; (JM) Departamento de Ecoloxia e Bioloxia Animal,
Facultade de Ciencias, Universidade de Vigo, E-36200, Vigo, Spain
Abstract.—Parapionosyllis cabezali, a new species of Exogoninae (Poly-
chaeta: Syllidae) from the Atlantic coast of Spain, is described. This new spe-
cies is characterised by the shape of the simple and compound setae, small
dorso-ventral gradation in the length of the blades, and by the presence along
the body of two types of parapodial glands, which open dorsally separately
from each other.
Campoy (1982) identified three specimens collected in Punta Endata, Gui-
puzcoa, NE Atlantic, Spain, as Parapionosyllis cf. gestans (Pierantoni, 1903).
Subsequently, during a study of the taxonomy and ecology of the sublittoral
soft bottom benthic polychaetous annelids in the Ria de Ferrol and Ria de
Baiona, both located on the Atlantic coast of NW Spain, more specimens of
this species were collected and reported by Parapar (1991), Parapar et al. (1994)
and Moreira (1999). Examination by SEM of several specimens collected in
the Ria de Baiona (NW Spain) has provided evidence that these organisms
represent a new species. The specimens are characterized by the shape and
dorso-ventral gradation in the length of the compound setal blades and by the
presence along the body of two types of parapodial glands. In this paper, we
describe and discuss this new species. The diagnosis of the genus was provided
by Fauvel (1923) and San Martin (1984).
Materials and Methods
The type specimens (holotype and para-
types) were collected by the junior author
(JM) in littoral soft bottom areas of the Ria
de Baiona (Galicia, NW Spain) at 9-12 m
depth. Additional specimens were collected
in Galicia, in the Ria de Ferrol. Samples
were taken both by means of a naturalist
rectangular dredge and a Van Veen dredge
in Dec 1995 in the case of the type series
and in 1987 and 1989 in the Ria de Ferrol.
Samples with the letters VD denote sam-
pling for the vertical distribution of animals
in the sediment. In this study, samples were
taken by scuba diving and by pushing a 50
cm corer into the sediment. Animals taken
from the sediment were fixed in 10% buff-
ered formalin and preserved in 70% etha-
nol. Body and setae measurements provided
in the description refer to the holotype, the
width was measured across the proventri-
culous and excludes cirri, parapodia and se-
tae. Observations, drawings, and measure-
ments were obtained using a microscope
with interference contrast optics (Nomar-
sky). Drawings were made with the aid of
a drawing tube. The SEM micrographs
were taken at the Servicio Interdepartamen-
tal de Investigacié6n of the Universidad Au-
tonoma de Madrid, Spain. The types are de-
VOLUME 113, NUMBER 2
posited in the Museo Nacional de Ciencias
Naturales de Madrid, Spain.
Family Syllidae Grube, 1850
Subfamily Exogoninae Rioja, 1925
Genus Parapionosyllis Fauvel, 1923
Parapionosyllis cabezali, new species
Figss 1,72
Parapionosyllis cf. gestans.—Campoy,
1982: 267-269, fig. 18.—Parapar et al.,
1994: 96-97, fig. 2.—Moreira, 1999:
319-323, fig. 26. [Not Parapionosyllis
gestans Pierantoni, 1903].
Material examined.—Ria de Baiona (Ga-
licia, NW Spain). Holotype and 23 para-
types. Sta. 8, Dec 1995, coarse sand, 12 m
(10 specimens); Sta. 9, Dec 1995, medium
sand, 10 m (10); Sta. 12, Dec 1995, coarse
sand, 9 m (4).
Additional material.—Ria de Ferrol (Ga-
licia, NW Spain), 39 specimens. Sta. 1, Jul
1987, muddy sand, 15 m (1 specimen). Sta.
3, Jun 1987, muddy sand, 11 m (1). Sta. 10,
May 1987, muddy sand, 18 m (2). Sta. 11,
Jul 1987, muddy sand, 20 m (1). Sta. 13,
May 1987, coarse sand, 14 m (2). Sta.
ISVD, Aug 1989, 20-25 cm in sediment
gem, 17 m (1). Sta. 17, Jul: 1987, very
eaarsersand, 25 mi (1). Stas, 22. Oct 1987,
very coarse sand, 20 m (1). Sta. 22VD, Aug
1989, O-—5 cm in sediment depth, 20 m (8).
Sta. 22VD, Aug 1989, 5—10 cm in sediment
depth (6). Sta. 22VD, Aug 1989, 10-15 cm
in sediment depth (1). Sta. 22VD, Aug
1989, 15—20 cm in sediment depth (2). Sta.
26, Aug 1987, very coarse sand, 15 m (5).
Sta. 37VD, Aug 1989, 0-5 cm in sediment
depth, 15 m (7). Ensenada de Baiona (Ga-
licia, NW Spain), 704 specimens. Sta. 8,
Dec 1995, coarse sand, 12 m (327 speci-
mens). Sta. 9, Dec 1995, medium sand, 10
mm (120). Sta. 12, Dec 1995, coarse sand, 9
m (236). Sta. 13, Dec 1995, medium sand,
8 m (18). Sta. 14, Dec 1995, medium sand,
9 m (3).
Description.—Body relatively long and
thin, with well marked segments. Holotype
S27
3.36 mm long, 0.24 mm wide with 39 seg-
ments. Prostomium semicircular to pentag-
onal (Figs. 1A, 2A); four eyes in an open
trapezoidal arrangement and two very small
anterior eyespots; eyes disappear soon after
fixation in alcohol. Antennae bottle or nine-
pin shaped, relatively long; median antenna
somewhat longer than the lateral ones, ap-
proximately of the same length as the pro-
stomium and palps together; lateral anten-
nae inserted ahead of anterior eyes; median
antenna between posterior eyes, near the
posterior margin of the prostomium. Palps
robust, equal to or slightly longer than the
prostomium, fused at bases but separated in
the anterior one-half; a slight junction su-
ture occasionally observed. Tentacular seg-
ment distinct but shorter and narrower than
the following segments; a pair of tentacular
cirri similar in shape to antennae but ap-
proximately one-half the length of the lat-
eral ones. Two ciliated nuchal organs (Fig.
2A, B) deeply imbedded between the pos-
terior margin of the prostomium and the an-
terior margin of the tentacular segment.
Anterior dorsal cirri short and oval, with
slightly enlarged bases (Figs. 1A, 2A); pro-
gressively more elongated and slender;
slightly longer than the parapodial lobes;
posterior dorsal cirri thinner and longer
than anterior ones (Fig. 1B). Two types of
parapodial glands (Fig. 1A, B); the smaller
ones beginning in the most anterior seg-
ments, including the first setiger, with gran-
ular content, and the larger ones with fi-
brillar material, located near the central
body line and starting from the proventric-
ular segments. These parapodial glands
open dorsally near the dorsal cirri, separate
from each other; each pore is provided with
a mobile, digitiform papilla which ejects the
products of these glands (Fig. 2C, D). Both
types of glands are dark red in colour. Para-
podial lobes short and conical, ending in a
small rounded papilla (Fig. 1C). Ventral cir-
ri digitiform, long and slender, shorter than
the parapodial lobes or similar in length.
Most anterior parapodia have about nine
compound setae; six or seven in the re-
528 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1. Parapionosyllis cabezali, new species. Holotype. A, anterior end and midbody, dorsal view; B,
posterior end, dorsal view; C, parapodium of midbody, posterior view; D, anterior dorsal simple seta; E, anterior
compound setae; EK aciculum; G, posterior dorsal simple seta; H, posterior compound setae; I, posterior ventral
simple seta. Scale bar a: 0.25 mm for A, B and 60 pm for C; scale bar b: 25 wm for D to I.
VOLUME 113, NUMBER 2
WN
nN
\O
t he
. iia
- 7 3 By
F 1 rt Wiebe hej
2 : r ~~ ?
° F ©
| a | a,
SU / 4 ij Sh
Magn r — ———s ere A aqn
Ox ,/ {/ 3000x
Fig. 2. Parapionosyllis cabezali, new species. SEM micrographs. A, anterior end, dorsal view; B, detail of
a nuchal organ opening; C, parapodial glands pores and papillae (arrows); D, detail of fibrilar material protruding
from a parapodial gland; E. long bladed, dorsal compound seta from anterior parapodium; EK mid-length and
short compound setae from midbody; G, posterior dorsal simple seta; H, posterior ventral simple seta.
530
maining anterior parapodia. Anterior dorsal
compound setae with thick shafts and het-
erogomph articulation (Fig. 1E); median
and ventral shafts with hemigomph articu-
lation (Fig. 2E, F). Shafts apparently
smooth but having a few subdistal spines.
Blades unidentate with spines on the mar-
gin and a longer sub-distal spine more ev-
ident on more dorsal and anterior setae.
Dorso-ventral gradation in blade length;
blades of the dorsal-most compound setae
about 25 ym in most anterior setigers, 20
im in the median ones and 15 pm in pos-
terior-most ones; with moderately long
spines on the margin; blades progressively
shorter towards the ventral part of the para-
podium (Fig. 1E, H), 10 wm in the ventral
most seta with shorter spines on the margin.
Solitary dorsal simple seta on each post-
proventricular parapodium, slightly thicker
than the shafts of compound setae, provided
with various spines on the margin, one of
which thicker and longer (Fig. 1D); most
posterior dorsal simple setae have fewer
spines on margin but are provided with a
larger subdistal spine (Figs. 1G, 2G). Soli-
tary ventral simple setae sigmoid, uniden-
tate, apparently smooth (Fig. 11) but pro-
vided with a few minute subdistal spines
(Fig. 2H). Solitary aciculum per parapodi-
um, relatively thick and with a rounded, en-
larged and apparently hollow tip (Fig. 1F).
Pharynx relatively long and thin, extend-
ing through five or six segments (Fig. 1A);
pharyngeal tooth located near the anterior
margin; anterior end of pharynx surrounded
by 10 papillae (Fig. 2A). Proventriculous
shorter than the pharynx, extending through
three or four segments, with about 22 rows
of muscle cells. Pygidium small, semicir-
cular, with two long and thin anal cirri, lon-
ger than the antennae (Fig. 1B). Mature
males provided with natatory capillary se-
tae and sperm packets in each segment be-
tween setigers 11 and 27; females carry
eggs ventrally.
Ecology.—Parapionosyllis cabezali, was
found in Punta Endata in sublittoral muddy
sand (70 m depth), in the Ria de Ferrol in
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
sublittoral muddy sand, coarse sand and
very coarse sand (11—25 m depth) with or-
ganic matter content between 0.12% and
0.20% and reaching 20 cm depth in sedi-
ment. In the Ria de Baiona this species was
found in sublittoral medium sand and
coarse sand (8—12 m depth).
Remarks.—Campoy (1982) and, more re-
cently, Parapar et al. (1994) reported this
species as Parapionosyllis cf. gestans (Pier-
antoni, 1903), due to its similarity to P. ges-
tans [as described by Fauvel (1923) and
Cognetti (1965)]. A more detailed study of
the specimens revealed two main differenc-
es between the species. P. cabezali shows
a moderate dorso-ventral gradation in the
length of the blades of the compound setae,
while in P. gestans all blades are similar in
length (Cognetti 1965). Unlike Campoy
(1982), we consider the setae illustrated to
be an accurate representation of the com-
plete set of setae in each parapodium. In the
same figure, the author provides detailed
sketches of the setae of three other species
(Parapionosyllis minuta, Parapionosyllis
elegans and Parapionosyllis labronica),
drawing the same number of setae for all of
them and showing the variation in the size
of the blades where it exists. Also, all spec-
imens of P. cabezali had numerous and
conspicuous parapodial glands, of two
kinds and different in size, which are not
described in the Mediterranean species. Per-
kins (1981) also report the presence of
paired parapodial glands in specimens of
Parapionosyllis longocirrata (Webster &
Benedict 1884) from the Atlantic coast of
the United States, but this species differs
from P. cabezali in that it has much larger
dorsal cirri, in the shape and spinulation of
the anterior dorsal simple setae and in the
length of the dorsal-most compound setae
blades (15 wm in P. longocirrata and 25
wm in P. cabezali). Parapionosyllis brevi-
cirra Day, 1954, a species also present in
the Atlantic coast of the Iberian Peninsula
(Parapar et al. 1994) and western Mediter-
ranean Sea, occasionally shows granular
and fibrillar inclusions in the dorsal part of
VOLUME 113, NUMBER 2
the parapodia (San Martin 1984), but this
well known species has much longer dorsal
compound setae blades (45 um) than P. ca-
bezali. The type-series of P. gestans (Pier-
antoni, 1903) was apparently lost, and it
was not possible to examine any type spec-
imens of this species. San Martin (1984)
recorded 12 described species of Parapion-
osyllis and provided a key for identification
in which P. gestans (Pierantoni, 1903) was
distinguished from P. cf. gestans of Cam-
poy (1982).
Distribution.—Parapionosyllis cabezali
seems to be distributed along the Iberian
Atlantic coast, from Basque Country (Cam-
poy 1982, Aguirrezabalaga 1984), Galicia
(NW Spain) in the Ria de Ferrol (Parapar
et al. 1994) and Ria de Baiona (Moreira
1999) to Punta Umbria (Huelva, SW Spain)
(Rodriguez & Viéitez 1992), although it has
not been reported in Portugal.
Etymology.—The species is named in
fond memory and honour of the late Luis
Cabezal Gomez, lecturer in the Escuela Pol-
itécnica Superior de Lugo (Universidade de
Santiago, Spain), brother-in-law of the first
author, in recognition of his love for nature
and friendship.
Acknowledgments
This paper was partially supported by the
projects “‘Aplicaciéns morfofuncionais da
quetotaxia de anélidos acuaticos de Gali-
cia” (XUGA10305B98) and ‘‘Fauna Ibéri-
ca IV”’ (PB950235) supported by the Xunta
de Galicia and Comision Interministerial de
Ciencia y Tecnologia respectively. The
drawings were made by J. Corbera, illustra-
tor for the project “Fauna Ibérica’’, and the
SEM micrographs were taken by E. Salva-
bse
dor of the Servicio Interdepartamental de
Investigacion (U.A.M., Spain).
Literature Cited
Aguirrezabalaga, F 1984. Contribuci6n al conocimien-
to de los Anélidos Poliquetos de la costa de
Guiptizcoa.—Munibe 36:119—130.
Campoy, A. 1982. Fauna de los Anélidos Poliquetos
de la Peninsula [bérica.—Publicaciones de Biol-
ogia de la Universidad de Navarra, Serie Zool-
6gica 7(1—2):1—781.
Cognetti, G. 1965. Sillidi mesopsammici delle secche
della Meloria (Livorno).—Archivio Zoologico
Italiano 50:65-—72.
Fauvel, P. 1923. Polychétes errantes. Faune de France,
5. Paul Lechevalier, Paris, 488 pp.
Moreira, J. 1999. Anélidos Poliquetos de sustratos
blandos infralitorales de la Ensenada de Baiona
(Galicia). Unpublished M.S. thesis, University
of Vigo, Vigo (Spain), 510 pp.
Parapar, J. 1991. Anélidos poliquetos benténicos de la
Ria de Ferrol (Galicia). Unpublished Ph.D. the-
sis, University of Santiago, Santiago de Com-
postela (Spain), 1104 pp.
, G. San Martin, C. Besteiro & V. Urgormi,
1994. Aspectos sistematicos y ecoldégicos de las
subfamilias Eusylline y Exogoninae (Polychae-
ta, Syllidae) en la Ria de Ferrol (Galicia, NO
Espana).—Boletin de la Real Sociedad Espa-
Nola de Historia Natural, Seccion Biologia
91(1-4):91-101.
Perkins, T. H. 1981. Syllidae (Polychaeta), principally
from Florida, with descriptions of a new genus
and twenty-one new species.—Proceedings of
the Biological Society of Washington 93:1080—
BLT:
Pierantoni, P. 1903. La gestazioni esterna.—Archivio
Zoologico 1(3—4):231—252.
Rodriguez, C. V. & J. M. Viéitez. 1992. Macrofauna
bentonica de los primeros metros del piso in-
fralitoral de la costa de Punta Umbria (Huel-
va).—Boletin del Instituto Espanol de Oceano-
grafia 8(2):327-338.
San Martin, G. 1984. Estudio biogeografico, faunistico
y sistematico de los poliquetos de la familia Sil-
idos (Syllidae: Polychaeta) en Baleares.—Uni-
versidad Complutense, Madrid, Tesis Doctora-
les, 187/84, 529 pp.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):532—541. 2000.
Paralarval gonatid squids (Cephalopoda: Oegopsida) from the
Mid-North Atlantic Ocean
Luisa I. Falcon, Michael Vecchione, and Clyde E E. Roper
(LIF) Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Autonoma de Mexico,
Apdo. Postal 70-305, D.E 045510, D.E, Mexico and National Museum of Natural History,
Research Training Program, Smithsonian Institution, Washington, D.C. 20560, U.S.A.
(MV) National Marine Fisheries Service, National Systematics Laboratory,
National Museum of Natural History, Washington, D.C. 20560-0153, U.S.A;
(CFER) Department of Invertebrate Zoology-Mollusks, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560-0118, U.S.A.
Abstract.—Ninety six gonatid cephalopod specimens (Oegopsida: Gonati-
dae) from the University of Amsterdam Mid-North Atlantic Plankton Expedi-
tions were analyzed and two species were identified: Gonatus steenstrupi (Kris-
tensen 1981) and Gonatus fabricii (Lichtenstein 1818). Gonatids were collected
only in spring and summer, despite sampling in autumn and winter. This paper
describes aspects of their development and reports their geographical distri-
bution in the central North Atlantic Ocean. Chromatophore patterns were the
most consistently useful characters for distinguishing between the species.
Among 34 measurements, Tentacle Length (TtL) relative to Dorsal Mantle
Length (ML) and number of suckers on Arms I-IV were useful for distinguish-
ing specimens >13 mm ML. Both species develop hooks from suckers on the
arms and tentacular clubs at ML >20 mm. Subtle differences were noted in
the morphology of the funnel pads except in the smallest specimens. Specimens
of G. steenstrupi >20 mm ML were collected at greater depths (250 to 995
m) than the smaller specimens (found at depths <200 m). Our data suggest
that 20 mm ML is the point of transition between paralarvae and juveniles of
G. steenstrupi, because specimens larger than 20 mm ML have well defined
hooks, and a juvenile vertical distribution is established.
Knowledge of early-life-history stages is
required for comprehensive understanding
of the ecology of squids. These stages pre-
sumably are most vulnerable to starvation
and predation and they occupy a separate
niche from older conspecifics (Vecchione
1987). Because of controversy about use of
the term “‘larva’’ for early life history stag-
es in cephalopods, Young & Harman (1988:
202) introduced the term “‘paralarva’’ for
‘‘a cephalopod of the first post-hatching
growth stage that is pelagic in near-surface
waters during the day and that has a dis-
tinctly different mode of life from that of
older conspecific individuals.’’ The paralar-
val concept includes both morphological
and ecological features, in contrast to the
definition of a “‘larva’’, which is based on
morphological differences from the adults.
The University of Amsterdam, Nether-
lands, conducted four research expeditions
in the North Atlantic Ocean between 55°N
and 24°N approximately along 30°W lon-
gitude. This research was designed to ‘“‘elu-
cidate the patterns of latitudinal diversity,
taxonomical variation below species level,
vertical variation and interaction of climate,
hydrographic features and ecology on mor-
phological variation of marine plankton”
(Van der Spoel 1981:1). The expeditions
VOLUME 113, NUMBER 2
were conducted during four consecutive
years (1980-1983), each during a different
season (Van der Spoel 1981, 1985; Van der
Spoel & Meerding 1983). Discrete-depth
samples were collected using opening/clos-
ing nets. Cephalopod paralarvae sorted from
the samples were donated to the National
Museum of Natural History, Smithsonian In-
stitution for systematic and ecological stud-
ies. Shea (1995) sorted the material into
families in preparation for subsequent stud-
ies. The present paper reports on paralarval
development and distribution of the squid
family Gonatidae in these samples.
The identification of paralarval cephalo-
pod stages is difficult because of insuffi-
cient collections which often come from in-
adequate sampling devices and methods,
and because of poorly understood taxono-
my, even in adults (Vecchione 1987). This
is true for the genus Gonatus in the North
Atlantic Ocean in which only one arctic/
boreal species was recognized, Gonatus fa-
bricii (Lichtenstein 1818), until Kristensen
(1981) described Gonatus steenstrupi from
boreal waters. At least five Gonatus species
are found in the North Pacific Ocean and
one in the Southern Ocean (Kristensen
1981). The primary morphological charac-
ters used to separate adults of the two North
Atlantic species are (a) the presence or ab-
sence of chromatophores on the ventral sur-
face of the head, (b) the shape of the funnel
organ, and (c) the patterns of hooks and
suckers on the tentacular clubs. The onset
of formation of hooks from suckers both on
the tentacular clubs and arms I-III seems a
good character to define the differences be-
tween paralarvae and juveniles in gonatids
(Young 1972, Kristensen 1977a). The pres-
ence of hooks presumably indicates a
change in feeding and therefore in the
squid’s role in the oceanic trophic structure.
The main goals of the present study were
to identify the species of paralarval gona-
tids in the Mid-North Atlantic collections,
to analyze early-life-history features that
could separate paralarvae from juveniles,
533
and to determine the distribution of the two
species in these samples.
Methods
Gonatid specimens were arranged by size
and their taxonomic identification began
with the largest specimens, then proceeded
sequentially to the smallest specimens. We
looked in particular for previously unrec-
ognized taxonomic characters for these par-
alarvae, in addition to using various taxo-
nomic guides (Kristensen 1981, Nesis 1987,
Roper et al. 1984). Fifteen specimens were
damaged and were excluded from some of
the quantitative analysis. Characters were
measured or counted on each of the 81 un-
damaged gonatids following Roper & Voss
(1983), including:
ML (Dorsal Mantle Length); MW (Mantle
Width); HL (Head Length); HW (Head
Width); ED (Eye Diameter); FL (Fin
Length); FW (Fin Width); TL (Total
Length); TtL (Tentacle Length); CL (Club
Length); AHI-IV (number of hooks on
Arms I-IV); ASI-IV (Arm I-IV Sucker
counts); ALI-IV (Arm I-IV Length); AWI-
IV (Arm I-IV Width).
We also measured the following characters:
D (Dactylus Length); M (Manus Length);
C (Carpus Length); CS (Club Sucker
Length); ForgL (Funnel Organ Length);
ForgW (Funnel Organ Width); ForgLl
(Funnel Organ Dorsal Pad Length);
ForgW1 (Funnel Organ Dorsal Pad Width).
Characters on damaged specimens were
measured when their condition permitted.
Three specimens of different sizes from
each of the two species were selected based
on condition, and their third arms and ten-
tacular clubs (left for G. fabricii and right
for G. steenstrupi, because of specimen
damage) were removed for scanning elec-
tron microscopical (SEM) analysis. The
specimens had been fixed in formalin and
preserved in 45% isopropanol. Tissue for
SEM was transferred through a dehydration
series to 100% ETOH prior to critical-point
drying, which was conducted using a Den-
534
ton Vacuum-1, Critical-Point Dryer. The
arms and tentacular clubs were examined
using a Leica 440 SEM to find morpholog-
ical features that could separate the species.
Results
In total, 96 specimens of the family Gon-
atidae were collected during the spring
(1980) and summer (1983) cruises. No gon-
atids were found in samples from the other
two cruises. The presence of two chromato-
phores on the ventral side of the head in G.
fabricii and their absence in G. steenstrupi,
the shape of the funnel organ and the de-
velopment and pattern of hooks and suckers
on the tentacular club were the basic fea-
tures that we used to distinguish these two
species. The 96 gonatids were separated
into 43 G. fabricii and 38 G. steenstrupi,
the remaining 15 specimens were too dam-
aged to determine species with certainty.
The 81 undamaged specimens ranged in
size from 1.6 to 31.6 mm ML in G. steen-
strupi, 3.3 to 24.1 mm in G. fabricii and
3.0 to 10.8 mm for the damaged specimens.
Gonatus fabricii paralarvae are charac-
terized by the presence of a pair of round
or oblong chromatophores on the ventral
surface of the head slightly anterior to the
ocular axis (see Kristensen 1981:67, fig. 3
for specimens larger than those reported
here). The dorsal pad of the funnel organ
in this species has an inverted V-shape with
very straight lateral sides (Fig. 1; cf. Kris-
tensen 1981:69, fig. 5). After the largest
specimen of G. fabricii (24.1 mm ML) was
identified, published taxonomic characters
could then be recognized in progressively
smaller specimens. The presence of chro-
matophores was the primary character used
to identify the smallest specimens of G. fa-
bricii. The funnel organ is so small in spec-
imens <3.6 mm ML that its shape can not
be determined confidently. Scanning elec-
tron microscopy on specimens of 24.1,
14.6, and 7.2 mm ML revealed that the pat-
terns of hooks and suckers on the tentacular
clubs of the largest G. fabricii were similar
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
to those described by Kristensen (1977a,
1981). The largest specimen (24.1 mm ML)
had one large hook, with three small hooks
and a sucker proximal to the large hook. A
concentration of small suckers occurs on
both dorsal and ventral sides of the club,
especially on the proximal end, where the
suckers form a large cluster (Fig. 2d).
The largest specimens of G. steenstrupi
(e.g., 31.6 mm ML) were identified follow-
ing the description of the holotype (Kris-
tensen 1981). The shape of the funnel organ
is characterized by a slight curve on the lat-
eral edges of the dorsal pad (Kristensen
1981: 83, fig. 20). This character can be
seen in our largest specimens (Fig. 3). The
absence of chromatophores on the ventral
surface of the head is the primary distin-
guishing character of the smallest G. steen-
strupi paralarvae available, which could not
be identified based on funnel-organ mor-
phology. Scanning electron microscopy was
used to examine specimens of 31.6, 19.2
and 9.6 mm ML. The tentacular club of the
largest specimen under SEM had the pat-
tern of hooks and suckers described by
Kristensen (1981) in which one large hook
is preceded proximally with four small
hooks and no suckers (Fig. 2c). At <20 mm
ML the hooks are not yet well developed
(Fig. 2a, b). Gonatus steenstrupi and G. fa-
bricii smaller than 20 mm ML have similar
sucker patterns in this central series on their
tentacular clubs.
The scatterplot of the relation between
TtL and ML for all the undamaged speci-
mens indicates a growth curve for G. steen-
strupi that diverges from that of G. fabricii
at ML >13 mm (Fig. 4). The relation be-
tween TtL and ML is more linear for G.
fabricii than for G. steenstrupi, perhaps be-
cause of the lack of large G. fabricii spec-
imens. However, all G. steenstrupi >13 mm
ML had tentacles that were longer than
those of similar-sized G. fabricii. Extension
of a quadratic function line fitted to the G.
fabricii data indicates that the difference in
TtL between species likely continues at
larger sizes, although a shift in growth pa-
VOLUME 113, NUMBER 2
Fig. 1. Growth series of G. fabricii. Scale bar = 1 mm.
Nn
eS)
OY
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
-e. 5
‘ ‘&
“ie RAL AS og an Oa ;
} bt
ee
rt
Fig. 2. Tentacular clubs. a—c) G. steenstrupi, 9.6, 19.2, and 31.6 mm ML respectively, d) G. Sabet 24.1
mm ML. Scale bars: a) 200 um, b) 300 wm, c) 1 mm, d) 1 mm.
rameters at the transition between paralar-_ ular clubs are found in specimens of both
vae and juveniles (Shea 1995) could either species larger than 20 mm ML. Kristensen
reduce or increase interspecific differences. (1981) described major differences between
Well-developed hooks on arms and ten- these species on specimens larger than 37
tacular clubs and well-differentiated tentac- mm gladius length, which is equivalent to
VOLUME 113, NUMBER 2
Fig. 3. Growth series of G. steenstrupi. Scale bar =
ML in gonatids. Our SEM analysis of arm
III suckers for G. steenstrupi of 31.6 mm
ML (Fig. 5a) and G. fabricii of 24.1 mm
ML (Fig. 5c) shows that no obvious differ-
ences occur, although the chitinous teeth of
y
1 mm.
the internal ring of the suckers are sharper
in G. steenstrupi. There is no difference be-
tween species in this character on smaller
specimens (Fig. 5b, d).
The vertical distribution of the paralar-
40
Bi Gonmaius steensirupt
@ Gonatus jabricti
€ 30
E
<
2
@ 20
&
oO
s
eae
2 40
0 %
0 10
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
20
Mantle Length (mm)
30 40
Dorsal Mantle Length (ML) vs. Tentacle Length (TtL). Quadratic function lines are fitted to the
scatterplots of the two species simply to identify trends in the data, rather than to test hypotheses of differences.
vae was analyzed for each cruise (Fig. 6).
During the spring cruise (1980), G. steen-
strupi larger than 20 mm ML were cap-
tured at the deepest sampling stations (250
to 995 m). The specimens smaller than 20
mm ML were collected in the upper 200
m, although one specimen of 2.8 mm ML
was caught between 390 and 510 m. Gon-
atus fabricii differed in spring vertical dis-
tribution, exhibiting greater variability in
depths for all sizes. All G. fabricii speci-
mens were captured shallower than 400 m
although samples were collected from
depths as great as 1750 m. The summer
cruise (1983) showed a similar distribution
for G. steenstrupi where specimens larger
than 20 mm ML were caught between 490
and 995 m and those smaller than 20 mm
ML were found in the upper 50 m. Gon-
atus fabricii were again found with greater
variability in their vertical distribution, as
specimens with 7.2 mm and 15.6 mm ML
were collected at the deepest stations
(1750 to 1000 m), while the rest of the
sampled population was found in the upper
100 m.
VOLUME 113, NUMBER 2
539
Fig. 3.
24.1 mm ML, d) G. fabricii, 14.6 mm ML. Scale bars: a) 20 wm, b) 20 wm, c) 30 wm, d) 10 pm.
Discussion
Van der Spoel (1981) reported that dur-
ing the 1980 spring cruise subarctic polar
water was present at depths greater than
500 m north of 50°N. The 1983 summer
cruise also may have sampled subarctic wa-
ter north of 53°N, as well as an isothermal
layer above the thermocline at depths great-
er than 90 m near 55°N. Both the spring
(1980) and summer (1983) cruises found a
well marked northern branch of the North
Atlantic Drift, although the southern branch
was much more marked in 1983 (Van der
Spoel 1985). The presence of Gonatus in
these samples likely resulted from the pres-
ence of cold subarctic water in the area.
Suckers in Arm III. a) G. steenstrupi, 31.6 mm ML, b) G. steenstrupi, 19.2 mm ML, c) G. fabricii,
Both Gonatus fabricii and Gonatus steen-
strupi, occurred in a previously unreported
distribution (54°53'54”"N 029°55'48"W to
48°58'54”"N 030°01'18”W). Earlier reports
(Kristensen 1977b, 1981: 62, fig. 1) listed
their distribution as nearer to the coast and
much more northerly, especially G. fabricii.
This new distribution extends the known oc-
currence of both species far offshore towards
the Central North Atlantic Ocean and more
southerly.
The change in vertical distribution by G.
steenstrupi larger than 20 mm ML and the
coincident presence above that size of well-
developed hooks on the arms and tentacular
clubs of both species may define the tran-
540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Mantle Length (mm)
1980 1983
a 0 10 20 30 40 b 0 10 20 30 40
Depth (m)
Gonatus steenstrupi
1500
Fig. 6. Depth of capture vs Dorsal Mantle Length (ML). a) G. fabricii in spring 1980, b) G. fabricii in
summer 1983, c) G. steenstrupi in spring 1980, b) G. steenstrupi in summer 1983. Diamonds indicate depth at
opening of net, squares depth at closing of net, vertical lines, depth range sampled.
sition between paralarvae and juveniles.
Differences between the species on speci-
mens larger than 20 mm ML include the
pattern of hook development from suckers
both on the tentacular clubs and the arms.
Additional specimens are needed to confirm
the apparent transition point between early-
life-history stages for G. steenstrupi and to
delineate such stages for G. fabricii. The
results reported here, however, do indicate
changes in both the morphological and eco-
logical characteristics at sizes about 20 mm
ML, particularly for G. steenstrupi.
Acknowledgments
The authors sincerely thank the Smith-
sonian Institution Women’s Committee,
which provided the funding for Luisa Fal-
con’s internship in the National Museum of
Natural History Research Training Pro-
gram. We are grateful to Guido Pastorino
for his help with the analysis of the data.
We acknowledge the SEM laboratory staff
at the NMNH, Walt Brown and Susann Bra-
den, for their assistance on our research.
Michael Sweeney, Division of Mollusks,
NMNH was of invaluable help. The authors
are grateful to the Research Training Pro-
gram Coordinator, Mary Sangrey, for mak-
ing this research possible. We appreciate the
careful reviews of an earlier draft by Uwe
Piatkowski and an annonymous reviewer.
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VOLUME 113, NUMBER 2
Roper, C. FE E., & G. L. Voss. 1983. Guidelines for
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ject 101 A).—Bulletin Zoologisch Museum,
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. 1985. List of discrete depth samples and open
net hauls of the Amsterdam Mid-North Atlantic
Plankton expeditions 1982 and 1983 (Project
101 A).—Bulletin Zoologisch Museum, Univ-
ersiteit van Amsterdam 10 (17):129-—152.
, & A. G. H. A. Meerding. 1983. List of dis-
crete depth samples and open net hauls of the
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):542—560. 2000.
Parabuccinum, a new genus of Magellanic buccinulid
(Gastropoda: Neogastropoda), with a description of a new species
M. G. Harasewych, Yuri I. Kantor, and Katrin Linse
(MGH) Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560-0118, U.S.A.;
(YIK) A. N. Severtzov Institute of Problems of Evolution, Russian Academy of Sciences,
Leninski Prospect 33, Moscow 117071, Russia;
(KL) Zoologisches Institut und Zoologisches Museum der Universitat Hamburg,
Martin-Luther-King-Platz 3, D-20146 Hamburg, Germany
Abstract.—A new genus, Parabuccinum, is established for four bathyal Ma-
gellanic species, of which one, Parabuccinum rauscherti, is described as new.
The remaining three species were originally described in the Antarctic genus
Chlanidota, from which they differ in the morphology of their protoconch,
operculum, radula, stomach, and male reproductive system. Parabuccinum ap-
pears to be most closely related to the monotypic circumAntarctic genus Neo-
buccinum, but may be distinguished on the basis of shell and penis morphology,
as well as by the size and shape of the gland of Leiblein. Parabuccinum is
endemic to the Magellanic Province, and is the first record of the subfamily
Bucinulinae in the malacofauna of this region.
The genus Chlanidota Martens, 1878 had
been considered to be one of the more di-
verse and wide-ranging members of the an-
tiboreal buccinoidean radiation. Dell (1990)
included 13 species within this genus, some
tentatively, among them four species that he
described as new. A more recent revision
(Harasewych & Kantor, 1999) divided the
genus Chlanidota into two subgenera,
Chlanidota sensu stricto, with five species,
and Chlanidota (Pfefferia) containing three
species. Five species were excluded from
Chlanidota on the basis of newly obtained
anatomical data as well as radular and oper-
cular morphology.
Three of the species described by Dell
(1990) (1.e., Chlanidota bisculpta, C. eltan-
ini, and C. polyspeira), all from the Magel-
lanic Province, differ substantially from all
species of Chlanidota. In this publication,
we propose a new genus to include these
Species, as well as an additional species
more recently collected by R/V Polarstern
and R/V Vidal Gomaz and described herein.
A review of each of the species included in
this new genus is provided.
Materials and Methods
This report is based primarily on material
collected by the United States Antarctic
Program (USAP), housed at the National
Museum of Natural History, Smithsonian
Institution (USNM), and consisting mostly
of the type material for the three species
described by Dell (1990). Additional sam-
ples, including alcohol preserved anatomi-
cal material, were collected at two stations
of the R/V Polarstern and one station of the
R/V Vidal Gomaz. These samples are main-
tained in the collections of the Zoologisches
Institut und Zoologisches Museum der
Universitat Hamburg (ZMH).
In the material examined sections, ““spec-
imen’’ denotes live collected material,
while ‘shell’? refers to records containing
only the dead collected shells.
Shell and operculum measurements were
obtained for representative specimens of
VOLUME 113, NUMBER 2
each species (n = 10, when available), as
detailed in Harasewych & Kantor (1999:
fig. 1). Anatomical descriptions are based
on gross dissections of preserved material.
Radulae were removed by dissection,
cleaned using diluted bleach (NaCI1Q), coat-
ed with carbon and gold, and examined us-
ing a Hitachi $570 Scanning Electron Mi-
croscope (SEM).
Photographs were obtained using a Leaf
Lumina Digital Scanning Camera. Optical
and SEM images were processed using
Photoshop Version 5.02 (Adobe).
The following abbreviations are used in
the text: SL—shell length, AL—aperture
length, BWL-—body whorl length, SW—
shell width.
Systematics
Order Neogastropoda Wenz, 1938
Superfamily Buccinoidea Rafinesque,
1815
Harasewych & Kantor (1999:256) pro-
vided a brief overview of the current state
of buccinoidean systematics and summa-
rized several of the more prevalent classi-
fications. Until the phylogenetic relation-
ships of the higher taxa of Buccinoidea are
reassessed on a global scale, we provision-
ally continue to use the family Buccinulidae
and its subdivisions, as defined by Powell
(1951), to include the many distinctive aus-
tral buccinoidean taxa.
Family Buccinulidae Finlay, 1928
According to Powell’s (1951:151) con-
cept of Buccinulidae, the chief characteris-
tics of the group are tricuspid rachidian
teeth and an operculum with a terminal or
subterminal nucleus. In our ongoing studies
on Buccinoidea, we were able to identify
several anatomical characters, among them
the prevalance of fused salivary glands, and
a simple stomach that lacks a posterior mix-
ing area, that will likely serve to distinguish
Buccinulidae from Buccinidae and other
buccinoidean families.
543
Subfamily Buccinulinae Finlay, 1928
Powell (1951) subdivided the family into
three subfamilies, the Cominellinae, Buc-
cinulinae and Prosiphiinae based on the
morphology of the lateral teeth of the rad-
ula. Cominellinae were characterized as
having bicuspid lateral teeth, Buccinulinae
were diagnosed by their tricuspid lateral
teeth, while Prosiphiinae were recognized
by their lateral teeth with a long basal pro-
jection. However, taxa included in Prosi-
phiinae vary considerably in their radular
morphology, suggesting that resolution on
a fine scale may be possible within this
group.
The radular teeth of the four species in-
cluded in the new genus Parabuccinum all
have tricuspid lateral teeth, indicating that
this genus is referable to the subfamily Buc-
cinulinae.
The geographic range of the subfamily
Buccinulinae spans Australia, New Zea-
land, Antarctica, and the eastern Pacific
coast as far north as California (based on
Powell’s (1951) inclusion of the genus Kel-
letia in Buccinulidae). Parabuccinum ex-
tends that range into the Magellanic Prov-
ince.
Genus Parabuccinum, new genus
Type species.—Chlanidota bisculpta
Dell, 1996.
Description.—Shell small for family,
reaching 10.2—16.4 mm, depending on spe-
cies. Shell relatively thick, solid, elongate
or elongate-ovate in outline, spire high
(about 0.33 to 0.5 SL). Spiral sculpture of
thin to prominent low cords. Axial sculp-
ture ranges from fine growth lines to prom-
inent, sigmoidal, weakly prosocline ribs,
depending on species. Aperture narrowly
oval. Siphonal notch broad, slightly re-
curved dorsally, abaxial margin may form
ridge along fasciole. Periostracum thin,
smooth or finely hirsute, covering most of
Shell. Operculum large (0.53—0.60 AL),
Ovate, paucispiral, with nucleus rotated
clockwise, nearly 180° along long axis of
544
operculum. Proboscis of moderate length.
Radular ribbon long, triserial. Rachidian
tooth with 3 large cusps (central cusp larg-
est) emerging from shallowly arched,
straight-sided basal plate. Lateral teeth with
3 cusps, outer cusp longest, middle cusp
shortest, closely adjacent to innermost cusp.
Salivary glands small, fused. Valve of Lei-
blein well defined. Gland of Leiblein very
small, tubular. Stomach broadly U-shaped,
without caecum. Penis long, with flattened
distal surface, bordered by thickened edge.
Papilla long, cylindrical, situated in middle
of distal surface, surrounded by very nar-
row circular fold at its base.
Remarks.—Conchologically, species of
Parabuccinum somewhat resemble juve-
nile specimens of Chlanidota, but differ in
having shells that are more slender and
have a large, invariably well preserved
protoconch, a relatively tall spire, and a
very thin periostracum. In contrast, the
protoconch of Chlanidota remains un-
known, despite the large number of spec-
imens (including juveniles <10 mm SL)
that were studied (Harasewych & Kantor,
1999). Anatomically, Parabuccinum dif-
fers from Chlanidota in having a propor-
tionally larger, paucispiral operculum, a
very long and thin-walled siphon, a very
small gland of Leiblein, a penis with a flat-
tened distal surface with a thickened edge,
and a stomach that overlies a portion of the
esophagus.
There are some conchological similari-
ties between this new genus and Buccinella
jucunda (Thiele, 1912), which differs in
having a prosiphiine radula (Thiele 1912:
pl. 16, fig. 13) and an operculum with a
terminal nucleus (pl. 13, fig. 20). Parabuc-
cinum, especially P. eltanini, also resembles
Neobuccinum eatoni (Smith, 1875) in gen-
eral shell outline, operculum and _ proto-
conch morphology.
Parabuccinum is endemic to the Magel-
lanic Province, and has a bathymetric range
of 247 to 2165 m.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Parabuccinum bisculptum (Dell, 1990)
Figs. 1, 2A—E, 3, 4, 7A—B, Table 1.
Chlanidota bisculpta Dell, 1990:185,
figs 291,294, 295,75 he.
Description.—Shell (Fig. 1) large for ge-
nus (to 14.8 mm), solid, white, elongate,
with angulated shoulder. Protoconch (Fig. 7
A-B) large (diameter 1.33—1.67 mm), of
about 2.25 smooth, glossy, convex, slightly
flattened whorls. Protoconch diameter/pro-
toconch height = 1.02—1.09. Transition
from protoconch to teleoconch well marked
by onset of axial and spiral sculpture. Te-
leoconch of up to 3.25 convex whorls. Su-
ture impressed. Spiral sculpture of raised,
distinct, narrow cords (12-16 on body
whorl, 5—7 on penultimate whorl), half the
width of intervening spaces. Axial sculp-
ture of raised, narrow, weakly sinuate, op-
isthocline ribs (18—22 on first teleoconch
whorl, 16—24 on subsequent whorls) that
form nodules at intersections with spiral
cords. Aperture large (0.51—0.65 SL), nar-
row, ovate, deflected from shell axis by 15—
19°. Outer lip, simple, rounded anteriorly,
shouldered posteriorly. Columella ~0.5 to
0.67 AL, weakly concave, with strong si-
phonal fold. Callus of thick glaze overlying
parietal region, siphonal fasciole. Siphonal
notch narrow, slightly dorsally recurved,
with nearly straight columellar and rounded
apertural margins that define borders of fas-
ciole. Ridge margin of fasciole formed by
apertural margin of siphonal notch. Shell
color chalky white, aperture weakly glazed.
Periostracum thin, transluscent, light yel-
lowish, thinly lallemose, with fine perios-
tracal hairs produced at intersections of spi-
ral threads with axial growth lines. Oper-
culum (Fig. 1E) medium-sized (0.53 AL),
elongate ovate, strongly coiled, with nucle-
us rotated clockwise, nearly 180° to long
axis of operculum.
Anatomical data based on single imma-
ture, male specimen (Fig. 1 I, R/V Polar-
stern, Sta. 10-109), from which only a por-
tion of the animal was recovered.
External anatomy.—(Fig. 2 A—B). Man-
VOLUME 113, NUMBER 2
Fig. 1. Parabuccinum bisculptum (Dell, 1990). A—C. Holotype, USNM 860128, off Burdwood Bank,
53°08'S, 59°23’'W, in 567-578 m (R/V Eltanin, Sta. 340). D—E. Paratype 1, USNM 860129, off Falkland (Mal-
vinas) Islands, 51°56’'S, 56°39’ W, 855-866 m (R/V Eltanin, Sta. 557). D. Apertural view of shell. E. Operculum
(coated with ammonium chloride). E Paratype 2, USNM 860129. Same locality as paratype 1. G—I. Off Cape
Horn, 55°44.0’S, 66°14.5’W, 430-397 m (R/V Polarstern, Sta. 40-109; G, H. ZMH, I. USNM 892150). Scale
bar = 5 mm for shells, 2 mm for operculum.
546
tle cavity spans just over 0.5 whorl, kidney
(Fig. 2A, k) % whorl. Upper whorls of vis-
ceral mass unknown. Columellar muscle
comprises 1.5 whorl, thick and narrow, at-
taching to shell at rear of mantle cavity.
Foot large, oval, broader anteriorly. Body
color very light yellow, nearly white, with-
out pigmentation. Head small, tentacles
(Fig. 2 A, B, ten) very long, narrow, widely
separated, gradually tapering. Eyes absent.
Siphon (Fig. 2 A, B, s) very long (0.75
AL), thin-walled, open.
Alimentary system.—Proboscis (Fig. 2
C—D, pr) of moderate length when contract-
ed (0.44 SL, 0.73 AL), smooth, unpig-
mented. Proboscis sheath thin-walled,
transparent. Mouth (Fig. 2 C, m) triangular
slit. Proboscis retractors (Fig. 2 C, D, prr)
broad, extremely thin, attached to middle
part of rhynchodaeum on left side, to its
base on right side. Buccal mass spans
~=().67 proboscis length. Radular ribbon
(Fig. 3) 1.55 mm long (0.39 AL), ~100—
140 pm wide (~0.015 SL, 0.025 AL), tris-
erial, consisting of about 60 rows of teeth.
Rachidian teeth with 3 closely spaced cusps
(central cusp slightly longer, wider than lat-
eral cusps) on posterior portion of basal
plate. Anterior margin of basal plate not
thickened, overlaid by adjacent tooth. Lat-
eral teeth usually with 3 cusps. Outer cusp
roughly twice as long, half as wide as inner
cusp. Intermediate cusp very thin, adjacent
to, equal in length to inner cusp. In one
specimen (Fig. 3 A—B) intermediate cusp
split into two separate cusps of equal size
along portion of radular ribbon. Salivary
glands (Fig. 2 D, sg) small, fused, situated
above nerve ring along left side of probos-
cis. Valve of Leiblein (Fig. 2 C, vL) well
defined, although not large, pyriform.
Gland of Leiblein (Fig. 2 C, D, gL) very
small, short, tubular, uncoiled, whitish,
opening into oesophagus without constric-
tion, just posterior to nerve ring (Fig. 2 C,
nr). Oesophagus narrow, thin-walled,
broader posteriorly. Stomach (Fig. 2 E, st)
broad, simple, U-shaped, without caecum.
Stomach simple, broad, overlies posterior-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
most oesophagus. Internal structures of
stomach not well-preserved.
Male reproductive system.—Specimen
immature male, with very short, dorso-ven-
trally flattened penis (Fig. 2B, p) lacking
defined papilla.
Type locality.—Off Burdwood Bank,
53°08'S, 59°23’'W, in 567-578 m (R/V El-
tanin, Sta. 340).
Type material.—Holotype (Fig. 1 A—C),
USNM 860128, from the type locality; Par-
atypes 1-6, USNM 860129, Paratype 7,
NMNZ ME .56615, off Falkland (Malvin-
as) Islands, 51°56’S, 56°39'W, 855-866 m
(R/V Eltanin, Sta. 557); Paratypes 8—9
(Paratype 9 is a specimen of Parabuccinum
rauscherti, new species, see below), USNM
860130, Paratype 10, NMNZ MEF .56616,
off Cape Horn, 56°06'S, 66°19’W, 384-349
m (R/V Eltanin, Sta. 740).
Material examined.—Type material in
USNM. 1 specimen + 1 shell, USNM
892150, 5 shells ZMH, off Cape Horn,
55°44.0'S, 66°14.5'W, 430-397 m, 16 May,
1996 (R/V Polarstern ANT XIII/4. Sta. 40—
109).
Distribution.—Off the Falkland (Malvin-
as) Islands and Cape Horn at depths ranging
from 349 to 866 m (Fig. 4).
Remarks.—This species most closely re-
sembles Parabuccinum polyspeirum and P.
rauscherti, new species. Parabuccinum bis-
culptum may be distinguished from P. poly-
speirum by its slightly smaller protoconch
(1.33—1.67 vs. 1.52—1.84 mm diameter), its
strongly shouldered rather than rounded
whorl profile, as well as by the presence of
fewer, stronger spiral cords and more pro-
nounced axial sculpture in the former (com-
pare Tables 1, 4). Characters that may be
used to differentiate P. bisculptum from P.
rauscherti, new species, are discussed in the
description of the new species.
Parabuccinum polyspeirum (Dell, 1990)
Figs. 4, 5, 7 E-E Table 2.
Chlanidota polyspeira Dell, 1990:186,
figs: 292.293.3113;
VOLUME 113, NUMBER 2 547
Fig. 2. Anatomy of Parabuccinum bisculptum (A-E, USNM 892150) and P. rauscherti (F—G, USNM
880616). A, B. Body removed from the shell. A. Right view. B. Anterior view of foot-head. C. Right lateral D.
Left lateral, views of anterior alimentary system (salivary glands removed to show valve of Leiblein in C). E.
Lateral view of stomach. K Mantle complex of organs (partially preserved, missing part is indicated by dotted
line). G. Anterior view of the body, mantle removed. Abbreviations: cme, cut mantle edge; ct, ctenidium; dg,
digestive gland; gL, gland of Leiblein; h, heart; k, kidney; m, mouth; nr, circumoesophageal nerve ring; oe,
oesophagus; op, operculum; os, osphradium; p, proboscis; poe, posterior oesophagus; prp, propodium; prr, pro-
boscis retractors; s, siphon; sd, salivary duct; sg, salivary gland; st, stomach; ten, cephalic tentacles; vL, valve
of Leiblein.
Description.—Shell (Fig. 5) large for ge-
nus (to 15.4 mm) solid, white, glossy, elon-
gate, with rounded shoulder. Protoconch
(Figs. 7 E-F) large (diameter 1.52—1.84
mm), consists of ~2.5 smooth, glossy, con-
vex, raised whorls. Protoconch diameter/
protoconch height = 1.12—1.28. Traces of
weak spiral cords may be present in final
0.25 whorl of protoconch. Transition from
protoconch to teleoconch abrupt, marked by
548
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1— Parabuccinum bisculptum (Dell, 1990). Measurements of shell characters. Linear measurements
in mm (nv = 9, including holotype).
Character Mean
Shell Length (SL) 9.9
Body Whorl Length (BWL) 76
Aperture Length (AL) 5.6
Shell Width (SW) 5.4
Protoconch diameter 1.53
BWL/SL 0.76
AL/SL 0.56
SW/SL 0.54
Number of axial ribs on Ist teleo- 19.5
conch whorl
Number of axial ribs on 2nd teleo- 19.7
conch whorl
Number of axial ribs on body 20.4
whorl
Number of spiral cords on body L357
whorl
Number of spiral cords on penul- 6
timate whorl
onset of pronounced spiral cords, followed
within 0.5 whorl by first, weak axial ribs.
Teleoconch of up to 3.25 convex whorls.
Suture impressed. Spiral sculpture of nu-
merous (19-26 on body whorl, 8—14 on
penultimate whorl) closely-spaced, raised
cords. Axial sculpture of raised, narrow,
sinuate, weakly prosocline ribs (16—18 on
first teleoconch whorl, increasing to 19-21
on body whorl) that form faintly cancellate
sculpture at intersection with spiral cords.
Aperture large (0.53—0.63 SL), narrow,
ovate, deflected from shell axis by 13-17°.
Outer lip thin, evenly rounded. Columella
=0.5 AL, weakly concave, with strong si-
phonal fold. Callus of thin glaze (very thin
in smaller specimens) overlying parietal re-
gion, siphonal fasciole. Siphonal notch nar-
row, slightly dorsally recurved, with nearly
straight columellar and rounded apertural
margins that form borders of fasciole. Ap-
ertural margin of siphonal notch demarcates
ridge margin of fasciole. Shell color chalky
white, aperture weakly glazed. Periostrac-
um very thin, smooth, translucent, tightly
adherent to shell surface. Operculum un-
known.
Type locality.—Patagonian Shelf, NE of
o Range Holotype
1.56 6.6-14.8 14.8
P33 5.0—11.6 11.6
1.08 4.0-8.9 8.9
0.83 3.9-8.2 8.2
0.12 1733—1-67 153
0.03 0.74—0.81 0.78
0.05 0.51—0.65 0.60
0.03 0.51—0.59 0.55
ies? 18—22 —
22535 16—24 19
3:26 16—24 2S
era 12-16 13
0.53 5-7 6
Islas de los Estados (Staten Island),
54°04'S, 63°35’W, in 247-293 m (R/V EI-
tanin, Sta. 369).
Type material.—Holotype (Figs. 5, A-—
C), USNM 860131, 3 paratypes, USNM
860132, 1 paratype NMNZ ME56617, all
from the type locality.
Material examined.—Type material in
USNM.
Distribution.—Known only from the
type locality (Fig. 4).
Remarks.—The species is known from
five shells. Dell (1990) regarded this taxon
to be very similar to P. bisculptum, but dis-
tinguished it on the basis of its narrower
Shell, larger protoconch, more numerous
spiral cords and axial sculpture that is
weaker, and becomes obsolete on the anter-
iormost part of the body whorl.
Parabuccinum polyspeirum is known
from a single station that represents the
shallowest record for any species of Para-
buccinum.
Parabuccinum eltanini (Dell, 1990)
Figs. 6,.8, 12, Table. 3.
Chlanidota eltanini Dell, 1990:184-5, figs.
290, 292, 297, 314.
VOLUME 113, NUMBER 2
i j a my
» of ter’: ie ‘ =
f “ wr ™ a
Gt Oe Sect . i» i 5 > oe =
BR Ss a See 4 ,
“a “ ; . ‘ \
Aish x we Sf”. Loe
fey " ; Se oe og
ns? A
oe
Radulae of Parabuccinum bisculptum (Dell, 1990). A. Dorsal, and B. right lateral (45°) views of the
central portion of the radular ribbon taken from animal in Fig. 11 (USNM 892150). C, D. Dorsal views of the
radulae of two specimens from R/V Polarstern, Sta. 40-109. Scale bars = 50 ym.
Fig. 3.
Description.—Shell (Fig. 6) largest of
genus (to 16.4 mm), solid, white, ovate,
with lower spire, rounded shoulder. Proto-
conch (Fig. 8) very large (diameter 2.06—
2.94 mm), dome-shaped, of about ~2.25
smooth, low, whorls. Protoconch diameter/
protoconch height = 1.30—1.52. Border be-
tween protoconch and teleoconch whorls
well demarcated, protoconch well pre-
served, teleoconch eroded. Teleoconch of
up to 2.75 broadly convex whorls. Suture
deeply impressed, with narrow, nearly flat-
tened rim. Spiral sculpture of numerous
(32-53 on body whorl, 17—24 on penulti-
mate whorl), adjacent, very low, fine, nar-
row, sometimes sinuous threads, covering
entire shell surface. Axial sculpture limited
to fine prosocline growth lines. Aperture
large (0.65 SL), ovate to broadly ovate, de-
flected from shell axis by 10—14°. Outer lip
simple, evenly rounded. Columella <0.5
AL, weakly concave, with broad siphonal
fold. Callus consisting of thick glaze nar-
rowly overlying parietal region, siphonal
550
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
200
Depth, m
500
Parabuccinum bisculptum (Dell, 1990)
“9 = type locality © = material examined
Parabuccinum polyspeirum (Dell, 1990)
+ = type locality
Fig. 4.
700
900
[234356 3
No. of specimens
Geographical distribution and bathymetric ranges of Parabuccinum bisculptum (Dell, 1990) and
Parabuccinum polyspeirum (Dell, 1990). Dashed line indicate 500 m isobath. The bathymetric range of P.
bisculptum is shown in black, that of P. polyspeirum in gray.
fasciole. Siphonal notch narrow, shallow,
slightly dorsally recurved, its margins de-
fining weak fasciole. Ridge margin of fas-
ciole runs from apertural margin of siphon-
al notch. Shell color chalky white, aperture
glazed. Periostracum thin, yellowish, tightly
adhering to the shell surface, bearing irreg-
ularly spaced very short hairs. Operculum
(Fig. 6H) large (0.60 AL), broadly ovate,
strongly coiled, with nucleus rotated clock-
wise, nearly 180° to long axis of operculum.
Radula illustrated by Dell (1990:fig. 297).
Type locality.—East of Falkland (Mal-
vinas) Islands, 51°58’S, 56°38’W, in 845—
646 m (R/V Eltanin, Sta. 558).
Type material.—Holotype (Fig. 6, A—C),
USNM 860124, paratypes 1-2, USNM
860125, paratype 3, NMNZ ME56613,
Table 2.—Parabuccinum polyspeirum (Dell, 1990). Measurements of shell characters. Linear measurements
in mm.
Character
Shell Length (SL)
Body Whorl Length (BWL)
Aperture Length (AL)
Shell Width (SW)
Protoconch diameter
BWL/SL
AL/SE
SW/SL
Number of axial ribs on Ist teleoconch whorl
Number of axial ribs on 2nd teleoconch whorl
Number of axial ribs on body whorl
Number of spiral cords on body whorl
Number of spiral cords on penultimate whorl
Holotype Paratype 1 Paratype 2 Paratype 3
JI) 92 15.4 oN 8.1
10.3 j bile fhe: 6.4
72 8.2 =a | 5.0
6s Fak 4.8 4.7
1.84 1.84 1.75 152
0.78 0.75 0.80 O79,
0.55 0.53 0.63 0.62
Ors2 0.50 0.53 0.58
16 16 17 18
22 18 24 2A
21 20 19 19
26 24 22 19
14 11 8 8
VOLUME 113, NUMBER 2
SAI
Nn
Fie: 5.
Patagonian Shelf, NE of Islas de los Estados (Staten Island), 54°04’S, 63°35'W, in 247-293 m D. Paratype 1.
E. Paratype 2. Both from type locality. Scale bar = 5 mm.
from the type locality. Paratypes 4—5 [par-
atype 5 (SL 4.8 mm) is not a buccinoidean,
but belongs in the family Cancellariidae
(Admetinae)] USNM 860126, E off Falk-
land (Malvinas) Islands, 51°56’S, 56°39’W,
855-866 m (R/V Eltanin, Sta. 557). Para-
Parabuccinum polyspeirum (Dell, 1990). A-—C. Holotype, USNM 860131, R/V Eltanin, Sta. 369,
types 6-7, USNM 860127, paratype 8,
NMNZ ME56614, E off Falkland (Malvin-
as) Islands, 54°09’S, 52°08’W, 419-483 m
(R/V Eltanin, Sta..1521):
Material examined.—Type material in
USNM.
Table 3.—Parabuccinum eltanini (Dell, 1990). Measurements of shell characters of intact specimens. Linear
measurements in mm. * = heavily eroded.
Character
Shell Length (SL)
Body Whorl Length (BWL)
Aperture Length (AL)
Shell Width (SW)
Protoconch diameter
BWL/SL
AL/SL
SW/SL
Number of spiral cords on body whorl
Number of spiral cords on penultimate whorl
Holotype Paratype Paratype
USNM USNM USNM
860124 860127 860125
135 16.4 15.8
113 13.6 13.0
8.8 10.3 =
8.8 10.0 9.6
2.06* 2.94 2.24*
0.84 0.83 0.80
0.65 0.63 0.63
0.65 0.61 0.61
BY 33 38
i 24 20
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 6.
E of Falkland (Malvinas) Islands, 54°09’S, 52°08'W, 419-483 m (R/V Eltanin, Sta. 1521). D. Apertural view.
E. Dorsal view of the protoconch. EF Lateral view of the protoconch showing transition between protoconch and
teleoconch whorls. G-H. Paratype, USNM 860125, type locality, E off Falkland (Malvinas) Islands, 51°58’S,
56°38'W, 845-646 m (R/V Eltanin, Sta. 558). G. Apertural view. H. Operculum (coated with ammonium chlo-
Parabuccinum eltanini (Dell, 1990). A-C. Holotype, USNM 860124. D—-F Paratype, USNM 860127,
ride). Scale bars = 1 cm for shells, 5 mm for protoconch, 2 mm for operculum.
Distribution.—Off Falkland (Malvinas)
Islands. Specimens collected at depths from
419 to 866 m. (Fig. 12).
Remarks.—This species differs from all
congeners in having a smoothly ovate shell
with a tall body whorl, a strongly prosoc-
line outer lip, an extremely large, dome-
shaped protoconch, and by the absence of
axial sculpture. Dell’s (1990:fig. 297) line
drawing of the radula of P. eltanini shows
a lateral tooth with a narrow basal plate and
inner cusp, which differs from those of oth-
er Parabuccinum (Figs. 3, 10) and some-
what resembles those of Chlanidota (see,
eg., Harasewych & Kantor, 1999 figs. 10 A,
C). Although we were unable to study the
VOLUME 113, NUMBER 2
anatomy of P. eltanini, we assign it to the
genus Parabuccinum on the basis of the
morphology of its shell, especially the char-
acteristic protoconch, and operculum.
Paratype 5, the smaller (SL = 4.8 mm)
of the two paratypes from R/V Eltanin sta-
tion 557 (USNM 860126), is not conspe-
cific with the rest of the type series, but is
a species referable to the cancellariid sub-
family Admetinae.
Parabuccinum rauscherti, new species
Figs. 2F—G, 7C—D, 9-12, Table 4.
Chlanidota bisculpta Dell, 1990 (par-
tim):185.
Description.—Shell (Fig. 9) small for ge-
nus (to 10.2 mm), solid, white, broadly
ovate to elongate, with flattened, shallow
subsutural rim, pronounced, rounded to an-
gulated shoulder. Protoconch (Fig. 7 C—D)
small (diameter 1.07—1.2 mm), of about
1.75 glossy, convex, inflated whorls. Pro-
toconch diameter/protoconch height =
1.10—1.31. Transition from protoconch to
teleoconch well marked by onset of spiral
followed immediately by axial sculpture.
First teleoconch whorl thinner than proto-
conch. Teleoconch of up to 3.25 convex
whorls. Suture strongly impressed. Spiral
sculpture of closely spaced cords varying in
strength from strongly pronounced to near-
ly smooth (16—40 on body whorl, 10—16 on
penultimate whorl), generally broader than
intervening spaces. Axial sculpture of
strong, broad, slightly sinuous, orthocline
ribs (13-17 on first teleoconch whorl, 15—
20 on subsequent whorls). Nodules formed
at intersections with spiral cords evident in
early whorls, abbraded on body whorl. Ap-
erture large (0.51—0.65 SL), narrow to mod-
erately ovate, deflected from shell axis by
12—15°. Outer lip evenly rounded, usually
slightly thickened. Columella <0.5 AL,
weakly concave, with strong, sharply de-
flected siphonal fold. Callus of thin to thick
glaze narrowly overlying parietal region
and siphonal fasciole. Siphonal notch mod-
erately broad, very slightly dorsally re-
ae
curved, with straight columellar, rounded
apertural margins defining borders of fas-
ciole. Ridge margin of fasciole formed by
apertural margin of siphonal notch. Shell
color chalky white, aperture weakly glazed.
Periostracum very thin, tightly adherant,
yellowish, weakly hirsute. Operculum (Fig.
9D, G) medium-sized (0.49—0.53 AL),
elongate ovate, strongly coiled, with nucle-
us rotated clockwise, nearly 180° to long
axis of operculum.
Anatomical data based on single, poorly
preserved male specimen (Fig. 9 E—E Par-
atype 1) from which only a portion of the
animal was recovered.
External anatomy.—(Fig. 2F—G). Foot
long (L/W ~2.5), oval, broad anteriorly, ta-
pering posteriorly, with thickened propo-
dium. Body color uniformly yellowish-
white. Head small short, tentacles (Fig. 3G,
ten) long, left longer than right, tapering
distally. Eyes absent. Mantle cavity (Fig.
3F) as wide as long. Mantle edge thickened,
with scalloped edge. Siphon (Fig. 3F) very
long (0.9 AL), thin walled, wide. Osphra-
dium ~0.67 mantle length, wide, with
curved axis. Ctenidium slighter longer, 0.67
as wide as osphradium, spans about 0.75 of
mantle length, formed of tall triangular la-
mellae that become narrower toward mantle
edge.
Alimentary system.—Proboscis of mod-
erate length in contracted position (0.35 SL,
0.6 AL), smooth, unpigmented. Proboscis
sheath very thin-walled, transparent. Mouth
opening triangular. Proboscis retractors
broad, extremely thin, asymmetrically at-
tached to proboscis sheath. Buccal mass oc-
cupies ~0.63 length of retracted proboscis.
Radula (Fig. 10) 1.54 mm long (0.33 AL),
~120-140 wm wide (~0.015-0.017 SL,
0.026—0.030 AL), of 65 rows of teeth, pos-
teriormost 3 nascent. Rachidian teeth tri-
cuspid (central cusp slightly longer, as wide
as lateral cusps). Lateral teeth usually with
3 cusps. Outer cusp nearly twice as long,
half as wide as inner cusp. Intermediate
cusp thin, equal in length to inner cusp. In
one specimen (Fig. 10 C) intermediate cusp
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
es
S205
Ss
Fig. 7. Protoconchs of Parabuccinum. A-B. P. bisculptum (Dell, 1990), ZMH. C—D. P. rauscherti, new
species. Paratype, ZMH 2813. E-E P. polyspeirum (Dell, 1990) Paratype in fig. 5 E. Arrows indicate transition
from protoconch to teleoconch. Scale bar = 500 wm for all figures.
VOLUME 113, NUMBER 2 Bi J
Table 4.—Parabuccinum rauscherti, new species. Measurements of shell characters. Linear measurements in
mm (n = 12, including types).
Paratype Paratype
Character Mean o Range Holotype ] 4
Shell Length (SL) 8.4 0.67 7.6—9.6 ist 8.1 8.1
Body Whorl Length (BWL) 6.5 0.63 5.4-7.4 6.1 6.3 6.2
Aperture Length (AL) 4.7 0.49 4.2-5.5 4.2 4.7 4.6
Shell Width (SW) 0 0.44 4.1-5.8 4.6 4.7 4.9
Protoconch diameter Nis 0.05 1.07—1.20 1.07 1.18 1.13
BWL/SL 0.76 0.03 0.71—0.80 0.79 0.78 0.77
AL/SL 0.56 0.05 0.51—0.65 0.55 0.58 0.57
SW/SL 0.60 0.04 0.54—0.67 0.59 0.58 0.60
Number of axial ribs on Ist teleoconch whorl 15.0 15 13-17 13 15 14
Number of axial ribs on 2nd teleoconch whorl 16.5 Ee 15-20 16 17 16
Number of axial ribs on body whorl 77 1.95 15-20 16 17 17
Number of spiral cords on body whorl 25.8 6.88 16—40 25 17 20
Number of spiral cords on penultimate whorl jaa | 2.28 10-16 12 10 10
fused with inner cusp along portion of rad-
ular ribbon. Valve of Leiblein well defined,
medium-sized, pyriform. Salivary ducts
join oesophagus just anterior to the valve.
Remainder of alimentary system poorly
preserved.
Male reproductive system.—Specimen
mature male, with long, broad penis (Fig.
11) extending length of mantle length. Dis-
tal end of penis transversely flattened, bor-
dered by thickened edge. Penial papilla
(Fig. 11B, pap) long, cylindrical, surround-
ed by narrow circular fold at the base.
Type locality.—Off Cape Horn, 55°44.0’S,
66°14.5'W, in 430-397 m. (R/V Polarstern,
ANT XIII/4. Sta. 40-109).
Type material.—Holotype (ZMH 2811),
paratypes 1—3 (USNM 880616), paratype 4
(ZMH 2812), paratypes 5—20 (ZMH 2813),
paratypes 21-30 (ZMH 28134), from the
type locality.
Material examined.—Type material. R/V
Eltanin. Sta. 339: Falkland (Malvinas) Is-
lands, Beauchene Island, 53°05’S, 59°31’'W,
512—586 m, 1 shell, USNM 870141; Sta.
740, off Cape Horn, 56°06’S, 66°19'W,
Fig. 8.
USNM 860127, E of Falkland (Malvinas) Islands, 54°09'S, 52°08’W, 419—483 m (R/V Eltanin, Sta. 1521). See
also Fig. 6 E-F Arrows indicate transition from protoconch to teleoconch. Scale bar = 1 mm.
Protoconch of Parabuccinum eltanini (Dell, 1990) at half the magnification of figure 7. Paratype,
556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 9. Parabuccinum rauscherti, new species. A-D. Holotype, ZMH 2811, R/V Polarstern ANT XIII/4,
Sta. 40-109, off Cape Horn, 55°44.0'S, 66°14.5’W, in 430-397 m. A-C. Shell. D. Operculum. E—G. Paratype
1, type locality, USNM 880616. E—-F Shell. G. Operculum. H. Paratype 4, ZMH 2812, type locality. I-J. R/V
Polarstern Sta. 40-114, off Cape Horn, 55°31.6'S, 65°56.8'W, 2165-2008 m, ZMH. Scale bar = 5 mm for
shells, 2 mm for operculum.
VOLUME 113, NUMBER 2
ro |
Fig. 10. Radula of Parabuccinum rauscherti, new species. A. Dorsal, and B. left lateral (45°) views of the
central portion of the radular ribbon taken from paratype 1, USNM 880616 (Figs. 9, E—F). C. Dorsal and D.
left lateral (45°) views of the central portion of the radular ribbon taken from paratype 4, ZMH 2812 (Figs. 9
G). Scale bars = 50 pm.
384-349 m, | shell (paratype of Chlanidota
bisculpta), USNM 860130. R/V Polarstern
ANT XIII/4: Sta. 40-114, off Cape Horn,
55°31.6'S, 65°56.8’W, 2165-2008 m, 18
May, 1996, ZMH, 29 shells. R/V Vidal
Gomaz: Sta. 42, Canal Concepcion,
50°35.7'S, 75°04.5'W, 250 m, 25 Oct 1996,
ZMH, 5 specimens.
Distribution.—Off the Falkland (Malvin-
as) Islands, Cape Horn and northward along
western coast of Chile to Canal Concep-
cion. Shells were trawled at depths ranging
from 349 to 2165 m. Live animals were col-
lected from 349 to 532 m (Fig. 12).
Etymology.—This new species is named
after Martin Rauschert who developed the
small dredge used to collect this species. Dr.
Rauschert is an amphipod taxonomist who
has worked for several years in the Antarc-
tic and Magellanic regions.
Remarks.—Parabuccinum rauscherti is
most similar in shell sculpture structure to
P. bisculptum from which it differs in hav-
ing a smaller, slightly broader shell (SW/SL
558
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 11.
Parabuccinum rauscherti, new species. Critical point dried penis of paratype 1, USNM 880616
(Figs. 9, E-F). A. Dorsal surface. Scale bar = 500 wm. B. Distal tip showing papilla. Scale bar = 200 pm.
=().60 in P. rauscherti, 0.54 in P. bisculp-
tum), a smaller protoconch (average diam-
eter = 1.16 mminP. rauscherti, 1.53 in P.
bisculptum) (compare Fig. 7A, B and 7C,
D). Parabuccinum rauscherti also has more
numerous and closely spaced spiral cords
(average = 25.9 on body whorl, 12.1 on
penultimate whorl) than P. bisculptum (av-
erage = 13.8 on body whorl, 6.0 on pen-
ultimate whorl) and less numerous (16.5 vs.
19.8) but wider axial ribs. Parabuccinum
rauscherti and P. bisculptum were taken to-
300
400
500
600
700
Parabuccinum eltanini (Dell, 1990)
@) = type locality
Parabuccinum rauscherti, new species
© =type locality
Fig. 12.
@ = material examined
@ = material examined
800
900
12:34
No. of specimens
B
2000
Depth, m
A
510° 20730
No. of specimens
Geographical distribution and bathymetric ranges of Parabuccinum eltanini (Dell, 1990) and Par-
abuccinum rauscherti, new species. Dashed line indicates 500 m isobath. A. Bathymetric range of P. rauscherti.
B. Bathymetric range of P. eltanini.
VOLUME 113, NUMBER 2
gether at two stations off Cape Horn (R/V
Polarstern Sta. 10-109 and R/V _ Eltanin
Sta. 740).
Parabuccinum rauscherti may be most
readily distinguished from either P. poly-
speirum or P. eltanini by its much smaller
protoconch and pronounced axial sculpture.
Parabuccinum rauscherti exhibits some
variation in shell outline and the number of
spiral cords.
The smaller of the two paratypes of
Chlanidota bisculpta, from R/V_ Eltanin.
Sta. 740 (USNM 860130, SL = 6.9 mm) is
actually a specimen of Parabuccinum raus-
cherti.
Discussion
Despite the limited availability of the an-
atomical data for Parabuccinum, it clearly
shares anatomical features with other Buc-
cinulinae, such as Chlanidota, Cavineptu-
nea and several yet to be named genera.
These similarities include proboscis mor-
phology, tricuspid rachidian and lateral rad-
ular teeth, a well defined valve of Leiblein,
fused salivary glands, a characteristic crop-
like structure of the posterior oesophagus,
and a broadly U-shaped stomach, lacking a
caecum (posterior mixing area). This stom-
ach morphology was found in all antiboreal
Buccinoidea studied thus far. Among north-
ern hemisphere buccinoideans, a stomach
lacking a caecum was recorded only in the
northern Atlantic buccinid Colus gracilis
(DaCosta, 1778) (see Smith 1967) and
deep-water Arctic species Mohnia (Tacita)
danielsseni (Friele, 1879) (see Lus 1981).
Parabuccinum has a number of unusual
features, among them a very long mantle
siphon, a very small, nearly vestigial gland
of Leiblein, and characteristic penis mor-
phology previously unrecorded in Bucci-
noidea.
With the exception of Parabuccinum, the
Magellanic buccinoidean fauna is com-
prised of genera attributed to the buccinulid
subfamily Cominellinae (e.g., Pareuthria,
Falsitromina, Parficulina). As there are no
559
closely related South American or Magel-
lanic Buccinoidea, Parabuccinum is likely
derived from the Antarctic Buccinulinae.
Powell (1951) attributed the following
Antarctic genera to the subfamily Buccin-
ulinae: Chlanidota, Pfefferia (a subgenus of
Chlanidota, as demonstrated by Harase-
wych & Kantor 1999), Neobuccinum, Prob-
uccinum, Cavineptunea and Bathydomus.
The differences between Parabuccinum
and Chlanidota have been discussed above.
Of the remaining Antarctic buccinuline
taxa, only one—the monotypic genus Neo-
buccinum, has a similar paucispiral oper-
culum. The shell morphology of Neobuc-
cinum eatoni (Smith, 1907) is similar to
that of Parabuccinum, especially to that of
P. eltanini. Both Neobuccinum and Para-
buccinum have large, dome-shaped proto-
conchs that appear to be more resistant to
errosion that their teleoconchs. The radula
of N. eatoni (illustrated by Numanami,
1996:fig. 94C) resembles that of P. eltanini
more that those of the other species of
Parabuccinum in that the central cusps of
the lateral teeth are larger and more distant
from the inner cusps. Parabuccinum and
Neobuccinum may be distinguished ana-
tomically, particularly on the basis of penis
morphology and the size and shape of gland
of Leiblein. The penis of Neobuccinum has
a broadly rounded distal end with a very
small, blunt seminal papilla, while the
gland of Leiblein in this species is large and
well developed (unpublished observations).
Neobuccinum eatoni has a circumantarc-
tic distribution extending along the Scotia
Arc, and to Kerguelen Island, as well as a
broad bathymetric range (5 to 2350 m). De-
spite the anatomical differences, we consid-
er Neobuccinum to be the most promising
candidate for sister group of Parabuccinum
among the presently known Antarctic buc-
cinoidean fauna. Parabuccinum thus rep-
resents an Antarctic component in the Ma-
gellanic malacofauna.
The distinctive paucispiral opercula of
Parabuccinum and Neobuccinum differ
from those of other austral Buccinoidea, yet
560
resemble opercula of the deep-water boreal
genus Mohnia (termed Mohnia type oper-
culum by Bouchet and Warén 1985:171).
Several species of Mohnia also have tricus-
pid rachidian and lateral teeth (Bouchet &
Warén 1985:178—-179) while at least one
species [Mohnia (Tacita) danielsseni, see
above] has a stomach lacking a caecum.
Mohnia is readily distinguished from Par-
abuccinum and Neobuccinum by numerous
conchological (e.g., presence of siphonal
canal) and radular (rachidian teeth with
squarish basal plate and lateral cusps lost or
closely juxtaposed to prominent central
cusp) characters. It 1s, as yet, unclear
whether these similarities are indicative of
a close relationship between these antipodal
taxa, or represent plesiomorphic characters
within the Buccinoidea.
Acknowledgments
We thank Dr. James McLean for his help-
ful comments on the manuscript. YIK
gratefully acknowledges Mrs. Mabs Mango
and Mr. and Mrs. Ed George for their kind
hospitality during the preparation of this
manuscript.
This research was supported in part by a
grant to YIK from the USAP Research Pro-
gram Award (Biological Collections from
Polar Regions at the National Museum of
National History) [National Science Foun-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
dation (Office of Polar Programs) Cooper-
ative Agreement OPP-9509761].
Literature Cited
Bouchet, P., & A. Warén. 1985. Revision of the north-
east Atlantic bathyal and abyssal Neogastropo-
da excluding Turridae (Mollusca, Gastro-
poda).—Bollettino Malacologico, Supplemento
I: 123-296.
Dell, R. K. 1964. Antarctic Mollusca.—Bulletin of the
Royal Society of New Zealand 27:1—311.
Harasewych, M. G., & Y. I. Kantor. 1999. A revision
of the Antarctic genus Chlanidota (Gastropoda:
Neogastropoda: Buccinulidae).—Proceedings
of the Biological Society of Washington 112:
253-302.
Lus, V. Y. 1981. On the abyssal species Sipho (Si-
Phonorbis) danielsseni (Friele) and Mohnia
mohni (Friele) (Gastropoda: Buccinidae).—Tru-
dy Instituta Okeanologii AN SSSR (Proceed-
ings of the Instute of Oceanology of the USSR
Ac. Sci.) 115:126—140 (in Russian).
Numanami, H. 1996. Taxonomic study on Antarctic
gastropods, collected by Japanese Antarctic re-
search expeditions.—Memoirs of National In-
stitute of Polar Research, Series E, Biology and
Medical Science 39:1—244.
Powell, A. W. B. 1951. Antarctic and SubAntarctic
Mollusca: Pelecypoda and Gastropoda.—Dis-
covery Reports 26:47—196, pls. 5—10.
Smith, E. H. 1967. The Neogastropod stomach, with
notes on the digestive diverticula and intes-
tine.—Transactions of the Royal Society of Ed-
inburgh, 67(2):23—42.
Thiele, J. 1912. Die antarktischen Schecken und Mus-
cheln.—Deutschen Sitidpolar-Expedition 1901—
1903 13:183-—285.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):561-571. 2000.
Porites arnaudi, a new species of stony coral
(Anthozoa: Scleractinia: Poritidae) from oceanic islands of the
eastern Pacific Ocean
Héctor Reyes-Bonilla and Juan P. Carricart-Ganivet
(HRB) Universidad Aut6noma de Baja California Sur, Departamento de Biologia Marina, Apdo.
Postal 19-B, La Paz, B.C.S. 23000, México; (JPCG) Departamento de Ecologia Acuatica,
ECOSUR, Apdo. Postal 424, Chetumal, Q. Roo, 77000, México
Abstract.—A new species of Porites (P. arnaudi) was found at depths of 7
to 37 m at Clipperton Atoll, France, and the Revillagigedo Islands, México,
oceanic islands located west of the American mainland (eastern Pacific). Col-
onies are uncommon but very distinctive because of the presence of tiered
plates, or laminae, expanding from a single base, smooth and concave surfaces,
and a lack of living tissue on the inferior part of the plates, except on its
growing edge. Calices are 0.8 to 1.4 mm in diameter, with wide walls. Cor-
allites have a free triplet with all three septa about the same size. Lateral septa
are fused in pairs and better developed than the rest. There are six to eight
pali, and one or two denticles per septum, and the color of tissue is greenish-
gray to dark brown. With this new species, the current total number of species
in the genus Porites in the eastern Pacific reaches nine, with most species
restricted to oceanic islands of the region.
In the last 20 years, taxonomic studies on
the scleractinian zooxanthellate fauna of the
eastern Pacific have noticeably increased in
number and quality (Wells 1983, Guzman
& Cortés 1993, Hodgson 1995) and as a
consequence, there is now fairly good
agreement on coral identities since some
key taxonomic problems have been re-
solved (Squires 1959, Wells 1983, Veron
1986). New research also has shown that
the actual richness of coral species from the
eastern Pacific is much higher than once
thought [e.g., originally less than 20 species
according to Veron (1993) and Paulay
(1997)]. For example, México, Costa Rica
and Panama have 20 to 25 species each and
almost 40 are known to inhabit western
America (Holst & Guzman 1993, Cortés &
Guzman 1998, Reyes-Bonilla & Lé6pez-
Pérez 1998). This perceived increase in
richness comes from new records for par-
ticular localities or areas (e.g., Reyes-Bon-
illa 1992, Cortés & Guzman 1998) and by
descriptions of new species (Budd & Guz-
man 1994, Glynn 1999).
Members of the genus Porites Link,
1807 are widely distributed in the eastern
Pacific, and they are one of the dominant
corals in this region, both in abundance and
species richness (Glynn 1997). To date,
eight species have been reported: P. aus-
traliensis Vaughan, 1918, P. baueri
Squires, 1959, P. lichen Dana, 1846, P. lob-
ata Dana, 1846, P. lutea Milne Edwards,
1860, P. panamensis Verrill, 1866, P. rus
Forskaal, 1775, and P. sverdrupi Durham,
1947 (Guzman & Cortés 1993, Glynn 1997,
Reyes-Bonilla 1999). Both P. panamensis
and P. lobata are present in coral commu-
nities from the Galapagos Islands, or the
Ecuadorian mainland, to México (Reyes-
Bonilla 1993, Glynn 1997), while P. sver-
drupi and P. baueri are endemic to the Gulf
of California and the Marias Islands, Méx-
ico, respectively (Squires 1959, Reyes-Bon-
illa & Lopez-Pérez 1998). Porites rus is an
562
Indo Pacific species that was reported from
Costa Rica in the early 1980’s, but its pop-
ulation may be extinct now (Glynn 1997,
Cortés & Guzman 1998). The remaining
three species are Indo Pacific also, but they
have been observed in the Revillagigedo
Archipelago, México, a group of four oce-
anic islands located about 300 km south-
west of the Baja California peninsula, or at
Clipperton Island, the only true atoll of the
eastern Pacific, 1300 km SW of the Mexi-
can mainland (Glynn et al. 1996, Ketchum
& Reyes-Bonilla 1997).
In this paper, a new species of Porites is
described. This coral is distinctive because
its colonies have been found exclusively in
the oceanic Clipperton and Revillagigedo
Islands, eastern Pacific. This finding in-
creases the known number of species of the
genus Porites to nine, a number even great-
er than that reported for the Atlantic-Carib-
bean area (Weil 1992, Veron 1993).
Abbreviations of repository institu-
tions.—USNM: United States National Mu-
seum of Natural History, Department of In-
vertebrate Zoology, Smithsonian Institu-
tion, Washington, D.C. 20560, U.S.A.;
MHNUABCS: Museo de Historia Natural
de la Universidad Aut6noma de Baja Cali-
fornia Sur. Departamento de Biologia Ma-
tina.«a; Paz .B:C:S3) 230803 Mexico:
ECOCHBC: Colecci6n de Corales de El
Colegio de la Frontera Sur, Grupo de Ecol-
ogia Bentonica. Apartado postal 424, Che-
tumal, Q. Roo, 77000, México. UMML:
University of Miami Marine Laboratory.
Rosenstiel School of Marine and Atmo-
spheric Sciences. 4600 Rickenbacker
Causeway, Miami, FL 33149, U.S.A.
Order Scleractinia Bourne, 1900
Suborder Fungiina Verrill, 1865
Superfamily Poritoidea Gray, 1842
Family Poritidae Gray, 1842
Genus Porites Link, 1807
Type species.—Porites polymorphus
Link, 1807 = Madrepora porites Pallas,
1766 (in part).
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Diagnosis.—Massive, ramose or encrust-
ing cerioid coralla. Corallites with calices
smaller than 2 mm in width, with two cy-
cles of septa. Septa commonly bear spines
and are formed by three or four trabeculae
that in their innermost portions are differ-
entiated as pali. A single trabecular colu-
mella, absent in certain species. Colony for-
mation by extratentacular budding. Coenos-
teum poorly developed (Wells 1956).
Distribution.—The genus is circumtrop-
ical, recorded in the Pacific, Indian and At-
lantic oceans, as well as in the Caribbean
Sea (Veron 1993). Fossil records are known
from the Cretaceous (?), Eocene to Recent,
but are more abundant and dominant in reef
communities since the Miocene (Wells
1956, Veron 1995). In the eastern Pacific
the genus is distributed from the Galapagos
Islands (O°N), Ecuador, to the northern Gulf
of California (31°N), México, including all
adjacent oceanic islands (Wells 1983,
Reyes-Bonilla 1993).
Porites arnaudi, new species
Figs. 1-6
Etymology.—Named after Gustavo Ar-
naud Vignon, the last Commander in Chief
of the Mexican Navy garrison on Clipper-
ton Island, who died tragically while on
duty on 1916.
Diagnosis.—(Terminology after Veron &
Pichon 1982 and Veron 1986) Oval shaped
colonies composed of tiered, thick plate-
like laminae (at least 10 mm in width). Cor-
allum originates from a single wide pedicel,
with smooth, concave and undulated sur-
faces, with no living tissue on the inferior
parts, except on its growing edges. Coral-
lites distinct, calices 0.8 to 1.4 mm wide,
with walls less than 1 mm wide. Septa rare-
ly bifurcated. Free triplet, although the ven-
tral directive and one lateral can be joined.
Lateral septa fused in pairs and better de-
veloped than the rest. One or two conspic-
uous denticles in the inner part of each sep-
tum. Six to eight pali which do not reach
the level of the wall; the palus of the triplet
VOLUME 113, NUMBER 2
563
Fis. 71;
b, side view. Both X0.49.
and dorsal directive less developed than
those of the lateral pairs. Two synapticular
rings, the external more defined. Living tis-
sue color greenish-gray to dark brown, with
pale margins.
Holotype.—USNM 100261 (Figs. la, b;
4a, b; 5a, b): length 213 mm, width 148
mm, height 111 mm, thickness of peripher-
Porites arnaudi, holotype, USNM 100261. Recent, 30 m depth, Clipperton Atoll, France. a, top view;
ical edge of folia 19 to 21 mm; collected
25 Nov 1997 by Juan P. Carricart-Ganivet.
Paratypes.—ECOCHBC 0107 (Fig. 2a,
b), MHNUABCS 1044, 1076, 1092, 1100,
1103 74581¢ (Pig. 3c; .d))- 1582: (Fig: Sa, b),
1583, UMML 8.1475 (Fig. 6a, b).
Type locality.—Northeast seaward slope
(sensu Glynn et al. 1996), off Clipperton
564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 2:
view; b, side view. Both X0.79.
Atoll, France (10°18’N, 109°13’W), at 30 m
depth.
Description.—Colonies submassive,
forming conspicuous tiered plates growing
outwards from a central base (discoidal
morphology; Figs. la, b; 2a, b); coralla can
attain more than 40 cm in diameter and
about 30 cm in height, being firmly at-
tached to the substrate. Smooth, concave,
and undulated upper surfaces, with thick fo-
Porites arnaudi, paratype, ECOCHBC 0107. Recent, 30 m depth, Clipperton Atoll, France. a, top
lia at least 10 mm in width (Fig. 3a—d). In-
ferior surfaces devoid of living tissue ex-
cept in their growing edges. Corallites 0.8
to 1.4 mm diameter, conspicuous, with
deep, polygonal calices (normally five to
six sides) having six to eight deep-set pal,
which do not reach the level of the wall.
Intercalicular distances 0.8 to 1.7 mm (Figs.
4a, b). Corallite walls less than 1 mm thick
and composed of three rows of denticles.
VOLUME 113, NUMBER 2 565
Fig. 3. Porites arnaudi, paratypes, top and side views of laminae from the edge of larger colonies: a, b,
MHNUABCS 1582; c, d, MHNUABCS 1581. Both recent, 21 m depth, Clipperton Atoll, France. All figures
x0.30.
566
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Porites arnaudi, variation of skeletal structure in the holotype USNM 100261. Notice diagnostic
characters such as polygonal calices and wide walls. a, X6; b, x9.
Twelve septa rarely bifurcated (Figs. 5a, b;
6a, b). The triplet normally has free mar-
gins, although it can also have the directive
and a lateral septum (generally that of the
right side) fused in the inner section. Lateral
septa arranged in four fused pairs, better de-
veloped and longer than the rest, and in its
innermost portion presents conspicuous pali
that rarely reach the level of the corallite
wall. Pali of the triplet and the dorsal di-
rective septa less developed than those of
lateral pairs, or may be absent (Fig. 6a, b).
One or two denticles per septum, which can
resemble small pali because of extensive
VOLUME 113, NUMBER 2
4
ae
Fig: 5.
567
Porites arnaudi, variation of skeletal structure in the holotype USNM 100261. Notice diagnostic
characters such as prominent pali, inconspicuous columella, one denticle per septum and free triplet (arrow). a,
X34; b, X66.
development and their position near the cal-
icular center. Columella quite variable,
sometimes appearing as a small column or
stylet, may be compressed dorsoventrally,
or may be absent. In some, a second prom-
inent protuberance about the same size as
the columella may be present near the inner
margin of the dorsal directive. Two synap-
ticular rings, the external more defined than
the inner palar. Tissue color from greenish-
gray to dark brown, with pale margins.
Distribution.—This species was first re-
corded by Glynn et al. (1996) at Clipperton
Atoll (southeast, northeast and southwest
568 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 6. Porites arnaudi, SEM photos of the paratype UMML 8.1475. Recent, 18 m depth, Clipperton Atoll,
France. a, group of calices X16. b, single calice X38.
sites) and at San Benedicto (19°18'N,
110°48’W) and Clarién islands (18°20’N,
114°44’W), Revillagigedo Archipelago, un-
der the name Porites sp. Later, Ketchum &
Reyes-Bonilla (1997) confirmed its pres-
ence in San Benedicto and Clarion, and also
at Socorro Island (18°45’N; 111°00’W),
calling it Porites sp. 1. However, no exact
locations of collection or observation sites
were presented in those papers. The holo-
type (USNM 100261, collected by J. P. Car-
ricart-Ganivet at 30 m depth, 25 Nov 1997)
and six other specimens on which the cur-
rent description is based (UMML 8.1475,
VOLUME 113, NUMBER 2
collected by P. W. Glynn at 18 m depth, 14—
26 Apr 1994; ECOCHBC 0107 collected by
A. Medina-Carcamo at 30 m depth, 25 Nov
1997; MHNUABCS 1581, 1582, 1583 col-
lected by H. Reyes-Bonilla at 21 m depth,
26 Nov 1997) were found in the ‘‘Poritid
zone” (sensu Glynn et al. 1996), at the
northeast seaward slope, Clipperton Atoll.
Other coralla of the same species were sam-
pled at Playa Norte and Punta Tosca (So-
corro Island; 18°47'25"N, 111°01'15’W,
MHNUABCS 1044, collected by J. EK Cas-
tellanos-Avila at 14 m depth, 30 Nov
1991), Bahia Azufre (Clarion Island;
18°20'47"N, 114°44'00"W, MHNUABCS
1092, 1100, 1103, collected by J. Ketchum
at 7, 7 and 10 m depth, 15 Dec 1994) and
Roca Trinidad (San Benedicto Island;
19°18'59"N, 110°49'00"W, MHNUABCS
1076, collected by H. Reyes-Bonilla at 15
m depth, 20 May 1995). As Porites arnaudi
has not been observed elsewhere in the cen-
tral Pacific or off American mainland, it is
considered to be endemic of the oceanic
Revillagigedos and Clipperton islands.
Ecology.—This newly described coral is
distributed at depths from 7 to 37 m, es-
pecially in steep rocky slopes, and often
overgrows adjacent coralla, mostly Porites
lobata and Pavona varians Verrill, 1864 in
Clipperton (Glynn et al. 1996) and P. lob-
ata and Psammocora superficialis Gardiner,
1898 in the Revillagigedos. It is not abun-
dant; Glynn et al. (1996) mentioned that
percent live cover was up to 7.4% in the
southeast side of Clipperton, while in the
Revillagigedos it is seldom found, although
it is locally abundant at Playa Norte (So-
corro Island), Bahia Azufre (Clarion Island)
and Roca Trinidad (San Benedicto Island).
Coralla commonly shows bite marks in the
outer edges of the plates, most likely pro-
duced by Arothron meleagris (Bloch &
Schneider, 1801) or Melichthys niger
(Bloch, 1786), both fishes that frequent lo-
calities where Porites arnaudi has been
found (Allen & Robertson 1994, Robertson
& Allen 1996). Bioeroders and serpulid
polychaetes were rare in colonies of this
569
species observed at Clipperton and practi-
cally absent in those collected for this pa-
per. In contrast, they were much more abun-
dant in the Revillagigedos, in particular the
bivalves Lithophaga spp., which commonly
bore the bases of the plates.
Comparison.—Because Porites arnaudi
has been found in Clipperton and at the Re-
villagigedo Archipelago, it must first be dif-
ferentiated from all other conspecifics pre-
sent in those localities: P. lobata, P. lutea,
P. lichen, P. australiensis and P. panamen-
sis (Ketchum & Reyes-Bonilla 1997,
Reyes-Bonilla 1999). The coral here de-
scribed is similar to P. lobata, as noted by
Glynn et al. (1996), because both species
have thick ledges with an inconspicuous
columella, well developed dorsal directive
septa and lateral pairs, palus of the dorsal
directive septum smaller than those of the
laterals, and septa of the triplet always free
(Veron & Pichon 1982, Veron 1986; Figs.
5a, b; 6a, b). Notwithstanding, the former
can be identified by its plate-like tiered col-
onies (Figs. la, b, 2a, b, 3a—d) which are
not massive or hemispherical as in P. lob-
ata. Also, it presents a well-defined syn-
apticular ring and different tissue color
(gray or brown in P. arnaudi and blue, pur-
ple or green in P. lobata). A third differ-
ence is that P. lobata has two denticles on
each septum, whereas P. arnaudi normally
has only one (Fig. 6b). Porites arnaudi and
P. lutea differ noticeably because the latter
is massive and has columniform lobes, shal-
low corallites and thin walls. In addition, in
P. lutea, pali reach the level of the walls,
all septa of the triplet are fused by a trans-
verse rod or in their margins (in a “‘tri-
dent’), bifurcate septa often appear and
columella is well developed. Porites arnau-
di can be easily separated from P. austral-
iensis because the latter has a large and
well-developed columella and pali that
reach the level of the wall denticles. Also,
in the triplet, the former species has lateral
septa about the same size as those of the
ventral directive (Figs. 5a, b, 6a, b), while
in P. australiensis the lateral septa are
570
smaller than those of the ventral directive,
and occasionally have fused triplets instead
of free ones. The morphology of the cor-
allum is quite different also, as P. austral-
iensis (like P. lobata) has massive, almost
spherical colonies (Veron & Pichon 1982,
Veron 1986). Porites panamensis can be
distinguished from P. arnaudi principally
because its colonies are very small (never
larger than 40 cm height and diameter), en-
crusting, massive or columnar, and its color
is bright green instead of gray or brown.
Other differences are that P. panamensis
has fewer pali (normally five) and thin
walls (Squires 1959). Porites lichen is an-
other species quite similar to P. arnaudi,
having six to eight pali and wide walls, one
denticle per septum, thick ledges around
colony bases, or appearing as laminar plates
(Veron & Pichon 1982, Veron 1986). Nev-
ertheless, they can be differentiated on the
basis of the corallites of P. lichen which
have septa of the triplet commonly fused in
their margins, while directive dorsal septa
of the same structure are shorter than lat-
erals. Also, coralla of this species appear as
columnar colonies, and when they occur as
laminae, these are thin and develop only in
the base of the coralla, a character never
presented in P. arnaudi (Figs. la, b, 2a, b,
3a—d).
Acknowledgments
Specimens for this work were collected
in a series of visits to the Revillagigedos
supported by UABCS, Comisi6n Nacional
para el Conocimiento y Uso de la Biodiv-
ersidad (contract FB342/H337/96), and
Baja Expeditions, and during the cruise
SURPACLIP-I to Clipperton and Socorro
islands, under the direction of Vivianne So-
lis Weiss (Instituto de Ciencias del Mar y
Limnologia, Universidad Nacional Auton-
oma de México, México City). James Ket-
chum, Juan Francisco-Castellanos, Alejan-
dro Granados, Miguel Angel-Garcia, Adri-
an Medina-Carcamo, Leonardo Ortiz and
Vivianne Solis collaborated in diving activ-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ities. Peter W. Glynn, Stephen D. Cairns,
James Ketchum and Andrés Lépez dis-
cussed and commented on preliminary ver-
sions of the manuscript. Special thanks to
Scott Monks for correcting the English, and
for his valuable comments on the first draft
of the manuscript. Photographs taken by
Humberto Bahena Basave at ECOSUR, and
scanning electron microscope images taken
at the USNM by Jarek Stolassky.
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
113(2):572-578. 2000.
A new species of Heterotella (Porifera: Hexactinellida: Euplectellidae)
from the West Indies
Henry M. Reiswig
Redpath Museum & Biology Department, McGill University, 859 Sherbrooke St. West, Montreal,
Quebec, Canada H3A 2K6, e-mail: cxhr@musica.mcgill.ca
Abstract.—Heterotella pomponae, a new species, is reported from near S.
Caicos Is., West Indies. This is only the second described species of the genus
and the first from the Atlantic Basin. Heterotella was erected by Gray to ac-
commodate Alcyoncellum corbicula Bowerbank from Isle de Bourbon (Re-
union), southwest Indian Ocean. The genus is distinguished from other Cor-
bitellinae by a microsclere which has controversial interpretation, as either a
oxyhexactin with long spines or an oxyhexaster with irregularly originating
terminal rays. The two species are distinguished by presence of sigmatocomes
in H. pomponae and their absence in H. corbicula.
‘The first taxonomically acceptable refer-
ence to a sponge now included in the genus
Heterotella was made by Bowerbank
(1867:358), where he clearly referred the
earlier published figure of a distinctive mi-
crosclere (Bowerbank 1858, 1864) to a
specimen labelled Alcyoncellum corbicula
from Isle de Bourbon in the Paris Museum.
The genus Alcyoncellum had a very con-
voluted early history, originally attributed
to a calcareous sponge and later considered
to be an uncertain synonym of the hexac-
tinellid genera Euplectella and Corbitella.
The status of many of these previously con-
founded generic names was clarified by
Gray (1967), where he erected the genus
Heterotella to contain only Bowerbank’s A.
corbicula. The two closely related genera,
Corbitella and Heterotella, were later re-
viewed by Ijima (1902), who described the
three then existing specimens of H. corbic-
ula from Isle de Bourbon (Reunion) in the
Museum National d’ Histoire Naturelle, Par-
is. The earlier figured and still diagnostic
microsclere of the genus was considered in
some detail. Ijima interpreted the oxy-
tipped spicule to be a hexactin with long,
irregular spines, but noted that this could
alternately be accepted as a oxyhexaster
with irregularly arising terminal rays. Kirk-
patrick (1910) reviewed the spicule in his
attempt to systematize spicule nomencla-
ture, and decided that it was an oxyhexaster
since the axial canal did not extend to the
ray tip. These distinctions remain arbitrary
since there is no method of objectively dif-
ferentiating spines from terminal rays.
No other Heterotella specimens have
been added to that type series collected
from Reunion in 1819 and 1857. Here I de-
scribe the first recent specimen of this ge-
nus, collected in 1994 from S. Caicos Is-
land, West Indies. The specimen represents
a new species of Heterotella and the first
member of the genus from a location other
than Reunion, Indian Ocean.
Materials and Methods
The specimen of an unidentified lyssa-
cine hexactinellid (the new species de-
scribed herein) was obtained on loan from
Harbor Branch Oceanographic Museum,
Fort Pierce, Florida (HBOM). The type se-
ries of H. corbicula was reviewed during a
1997 visit to the Museum National
d’ Histoire Naturelle, Paris (MNHN). Frag-
ments of two of the MNHN specimens were
VOLUME 113, NUMBER 2
returned to Montreal where skeletal prepa-
rations were made. Small pieces of dermal
and gastral surfaces were either whole-
mounted in balsam for light microscopy
(LM) or digested in hot nitric acid. Large
spicules in the resulting spicule suspensions
were rinsed, spread on microscope slides
and mounted in balsam. Smaller spicules
were dispersed on 25 mm diameter, 0.2 ~m
pore-size, nitrocellulose filters by filtration;
the filters were rinsed, dried and mounted
in balsam. Spicules were measured by com-
puter via a microscope-coupled digitizer.
Data are reported as: mean = St. deV. number
(range). Spicule drawings were prepared
from video-captured microscope images
imported into a computer drawing program
and traced on-screen. Samples for scanning
electron microscopy (SEM) were nitric-
acid-cleaned and mounted on stubs with ep-
oxy. Following gold-palladium coating,
specimens were viewed and photographed
with a JEOL JSM-840 SEM.
Systematics
Subphylum Symplasma Reiswig &
Mackie, 1983
Class Hexactinellida Schmidt, 1870
Subclass Hexasterophora Schulze, 1886
Order Lyssacinosa Zittel, 1877
Family Euplectellidae Gray, 1867
Subfamily Corbitellinae Ijima, 1902
Genus Heterotella Gray, 1867
Type species.—Heterotella corbicula
(Bowerbank, 1867).
Diagnosis.—(from Tjima 1902:32 for H.
corbicula, emended here). Corbitellinae of
saccular shape, the lateral walls constricting
distally to the margins of a terminal sieve-
plate. With numerous, round, irregularly ar-
ranged parietal oscula. Skeletal beams un-
fused or totally fused by synapticula. Prin-
cipalia parenchymalia as diactins; accesso-
ria as diactins and hexactins. With
distinctive microxyhexasters in which the
slender terminal rays are irregularly undu-
lating and do not all originate from a single
point (often interpreted as microxyhexac-
eH eo.
tins with irregular spines). Floricome and
graphiocome present, with or without sig-
matocome; discohexaster lacking.
Remarks.—A holotype has not been des-
ignated for H. corbicula from among the
original three specimens in the collections
at MNHN. A lectotype is here designated
as MNHN HX 23, Ijima’s specimen A, the
specimen upon which he based his exten-
sive spicule description. The specimen can
be identified as consisting of only the top
portion of a sponge from Isle de Bourbon
donated by Mr. Leschinault, 1819. The two
remaining specimens, MNHN HX 24 & 25,
Ijima’s B and C, thus become paralecto-
types. Lectotype designations are made
here with the purpose of clarifying the ap-
plication of the name to a taxon.
Heterotella pomponae, new species
Figs. 1-10, Table 1
Material examined.—Holotype: HBOM
0027-00019? km: -S) “ofes. Caicos’ IJs.,
21°28.84'N, 71°29.985'W, 467 m; col. A.
Wright via RMS Johnson SeaLink I, dive
3788, 31 Oct 1994; in alcohol.
Shape.—Entire specimen with basal disc
retrieved; broken during handling. Thin-
wall, sac-shaped sponge (Fig. 1), 13 cm tall
by 5.5 cm diam, attached to hard substrata
by short basal disk supporting a short, rigid
basal cup. Wall, 2.3 mm in maximum thick-
ness, perforated by numerous primary pa-
rietal oscula passing directly through the
wall, diameter 1.7 + 0.3,; mm (1.1-2.1
mm), spaced at 6.4 + 1.4,; mm (3.1—9.0
mm); smaller openings passing obliquely or
tortuously through the wall (secondary pa-
rietal oscula) have diameter on the dermal
side of 1.5 + 0.73, mm (0.4-—2.7 mm, Fig.
2) and on the atrial side 1.0 + 0.3,, mm
(0.5—2.0 mm, Fig. 3). Parietal oscula are ar-
ranged without apparent pattern. The ter-
minal osculum, 2.3 cm diam is covered by
a sieve plate and bordered by a vertical
marginal collar.
Surface texture.—Dermal tissue is deli-
cate and subject to detachment by gentle
574
Fig. 1. Heterotella pomponae n. sp. (holotype)
cut-away diagram reconstructed from photographs,
showing distribution of main parietal oscula, terminal
sieve plate and location of sinuous diactins in the base
of the atrium (wavy filled area).
water currents; gastral tissue is firmer and
more adherent to the skeletal framework.
Both surfaces are smooth to eye but irreg-
ularly lumpy under a dissecting micro-
scope. Prostalia are lacking; the fine tips of
sword hexactins projecting from the cush-
ion-like white tissues can be perceived only
with aid of a microscope; no regular grid
arrangement is detectable on either surface.
Living tissues with spicules are not present
in the sieve plate or marginal collar; mar-
ginalia absent, but may have existed in
younger stage.
Color.—White in life and preserved in
ethanol.
Skeleton.—The entire framework of the
holotype (basal cup, lateral walls, sieve
plate) has been solidified into a rigid struc-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ture by siliceous synapticulae so profuse as
to obscure the nature of the primary frame-
work-forming megascleres (Figs. 4-8).
These are presumed to be diactins since
hexactin nodes have not been found in this
structure and diactins are known to be prin-
cipalia of H. corbicula. Major siliceous el-
ements of most of the framework appear
randomly oriented (Fig. 4), but the outer
layer contains major longitudinal bundles
and the inner-most atrial bundles are trans-
versely oriented. The outer-most strands of
the framework are nodulated by secondary
silicification (Fig. 5) while the middle and
internal strands are smooth. Thickness of
the framework is 1.6 + 0.2,, mm (1.2—1.9
mm). In the basal region of the body (Fig.
1), a dense matt of sinuous diactins joined
by very short synapticulae or point cemen-
tation, is deposited on the internal surface
of the framework (Fig. 6); this matt nearly
fills the mesh spaces (Fig. 7), occludes
some parietal oscula and strengthens the
basal framework.
Elements of the sieve plate and marginal
collar are rigidified and nearly filled to solid
sheets by dense synapticulation (Figs. 8, 9).
The marginal collar extends up from the
body surface 1.9 + 0.4,, mm (1.4—2.4 mm),
as a nearly solid sheet of fused, transverse-
ly-oriented, siliceous spicules. Major
strands of the body framework extend up
the inner side of the collar and project out
into the oscular plane as sieve beams (Fig.
8). The sieve beams are unusual in their
vertical sheet-like form, being taller than
wide (septa-like), with beam height: 0.78 +
0.4,, mm (0.2-1.9 mm), width: 0.25 +
0.10,; mm (0.11—0.53 mm). Sieve pores are
variable in shape and size (Fig. 9), but are
mainly subtriangular with sides 1.5 + 0.4,,
mm (0.6—2.3 mm).
Loose spicules.—Diagrams of spicule
form and dimensions are summarized in
Figure 10 and Table 1.
Megascleres: The major surface spicules
of both dermal and gastral surfaces are
smooth, sword-shaped hexactins with the
short ray projecting from the tissue surface
VOLUME 113, NUMBER 2 +75
vine ;. 4
> =>
- @
“a th ‘
i, A ner +
Figs. 2-9. Heterotella pomponae n. sp. (holotype). 2-3. Photographs of dermal (2) and gastral (3) surfaces
with tissues, showing differences in aperture distributions; scale equals 1 cm (LM). 4—5. External surface of
acid-cleaned, fused framework of upper body wall; scale equals 1 mm (SEM). 6. External surface of basal region
with meshes filled with sinuous diactins; scale equals 1 mm (SEM). 7. Internal surface of the cup base covered
with dense pad of sinuous diactins; scale equals 1 mm (SEM). 8. Oblique vie of a fragment of the fused marginal
collar and sieve plate showing continuity of sieve beams with wall bundles: scale equals 1 mm (LM). 9. Side
view of a major sieve plate beam, top surface to left, showing complete fusion of constituent diactins; scale 0.1
mm (LM).
576
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 10. Loose spicules of Heterotella pomponae n. sp. (holotype). A. Superficial dermal and gastral sword
hexactins grading to pentactin. B. Choanosomal regular hexactins. C. Centrotylote diactin. D. Ends of sinuous
diactins. E. Oxyhexaster. F Floricome. G. Sigmatocome. H. Graphiocome centrum with long and short terminals
(raphides). Scale bars: Left 100 1m bar applies to A, B; right 100 wm bar applies to C, D; 25 wm bar applies
to E—-H.
(Fig. 10A); a moderate number of pentac-
tins (11%) may result by reduction of the
distal ray. These occur in mixtures of thin
and thick forms on both surfaces. True mar-
ginalia were not found—the marginal collar
was devoid of tissues or loose spiculation.
Nearly equal-rayed, sparsely-spined hexac-
tins (Fig. 1OB) occur throughout the choan-
osome in small numbers; they vary greatly
in size and may consist of two classes, but
frequency analysis was not performed.
Sparsely-spined, centrotylote diactins with
acute tips (Fig. 10C) are abundant through-
out the choanosome. Smooth, sinuous diac-
tins with rounded tips (Fig. 10D) pack the
lower framework. They occur as fragments
up to 6+ mm long in loose spicule prepa-
rations, presumably broken from‘their syn-
apticular connections; loose complete spic-
ules of this class have not been found.
Microscleres: All of the following mi-
croscleres, with exception of the short-
rayed graphiocome noted below, are found
in samples from both surfaces and the
choanosome. The most common micro-
sclere is a smooth, sharp-tipped form (Fig.
10E) which can be regarded either as an
oxyhexaster with very irregular secondary
VOLUME 113, NUMBER 2 577
Table 1.—Spicule dimensions of Heterotella pomponae holotype in zm; SD = standard deviation; n = number
of measurements.
Spicule type Length* + SD Range N Width + SD Range n
Hexactin, thick sword
distal ray 43° .24 8-101 SO 3 Se Ae «oa
tangential ray Se. > 15 52-121 a0, SS) 2 Cl a5 3-o48- 25
proximal ray 507, ='96 335-730 a0 Dy ae BD
Hexactin, thin sword
distal ray 90.25.17 35-118 50) (SOE OF VY 256 25
tangential ray 106 + 18 67-141 50): SScteOGul 2788.) 25
proximal ray 306.2 77 171-541 506 350 oP Oo 25
Hexactin, regular ray Ist 281 64-318 50, 321. = aS. 2s 30
Diactin, centrotylote S69) 257 320-1963 SUT yay 2 Eee. 3.8-12.3 50
Diactin, sinuous — — —=/9 23.1 2'6.3'>' 9.1=40:2 - “50
Oxyhexaster radius 50:5 + 6.8 B73-7hid 106. 20:04 Lt —eehy «25
oxyhexaster secondary ray LSB. 45 5.8—28.7 100 120.3 OBS: 0 Zo
Floricome radius Feel ae ce a 21644.6 357 54+1.0° 3474 25
floricome centrum radius Ned = O8 5.49.0 100. ADE O83 lhieasS =o
Sigmatocome radius |W ge Pike es 133—20'5) 6 100 0:6: 2502 O20 25
sigmatocome centrum radius 6.8 + 0.8 5.1-9.5 100 — Ed 0.37 9106-16 (25
Graphiocome centrum radius 82,209 6.3-11.1 IDO») (13:2 O3"°OS-19 25
graphiocome long secondary ray 86 + 9 63-151 100° * 0.92.02 .01641:2"'" 25
graphiocome short secondary ray 14.9 + 1.8 BIG —2-Or 100: 6) O6:2e10sike 04-09 125
@ Radius for microscleres, ° Width of terminal flange.
rays (preferred here) or an oxyhexactin with
long spines. The secondary rays (or spines)
number 1—4, most commonly 3, and ema-
nate from an elongate swelling at their junc-
tion. About 10% of these are without sec-
ondary rays but bear the swelling seen in
the astrose form. A typical floricome (Fig.
10F) with usually 7 (6—9) terminal rays,
bearing 3—5 claws on the terminal flange, is
common. Although these occur in a wide
size range, frequency analysis indicates
these compose a single class. An uncom-
monly occurring sigmatocome (Fig. 10G)
bears 25—35 sigmoid terminals, uniform in
length, from a single marginal whorl on
each recurved primary capitulum. Graphio-
come centra, with stumps of 30—50 terminal
attachments scattered across the face of
each discoid capitulum, (Fig. 10H) are un-
common. The presumed raphide-like, long
terminal rays are common on filters from
all tissues, but the short versions are found
only in gastral tissues. Intact graphiocomes
have not been encountered.
Etymology.—This species is named to
honour Dr. Shirley Pomponi in recognition
of her many years of invaluable contribu-
tions to the knowledge of tropical marine
biodiversity.
Remarks.—The vary large number of
shared characters between the new species
and H. corbicula leaves little doubt that the
new the West Indian form belongs to Het-
erotella. Most of the differences between
the two species can be attributed to their
different stages of maturation. Only speci-
men B of H. corbicula exhibits incipient
spicule fusion in its lower portion, while the
H. pomponae specimen has completely
fused its lyssacine net into a rigid frame-
work. The absence of free large choanoso-
mal diactins and marginalia in H. pompon-
ae are attributable to this terminal growth
stage, where further body extension
(growth) has ceased. Surface megascleres
and microscleres are almost identical in
form and size in both species. The dermalia
are thicker (to 30 wm) and the accessory
diactins longer (to 5 mm) in H. corbicula
than in H. pomponae (to 6.7 wm and 2 mm
578
respectively). The distinctive oxyhexasters
(oxyhexactins) are virtually identical in the
two species. The most unambiguous differ-
ence between the two is the presence of sig-
matocomes in H. pomponae and their ab-
sence in A. corbicula, confirmed by ex-
amination of new spicule preparations of
Ijima’s specimen A (the lectotype) and B
(the nearly intact paralectotype) using filter
techniques. Photographic records at HBOM
suggest that H. pomponae is moderately
common in the West Indian area. Younger
specimens with incompletely fused skeletal
networks are expected to be found in future
collections.
Acknowledgments
I thank C. Adams for bringing the spec-
imen to my attention, S. Pomponi and J.
Reed for providing access to the specimen,
photographs and data sheets, M. Kelly for
providing transparencies and motivation, C.
Lévi and K. Tabachnick for assisting in re-
view of the Paris material, and the latter for
his valuable comments on improving the
manuscript. This study was supported by an
operating grant from the Natural Sciences
and Engineering Research Council of Can-
ada.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Literature Cited
Bowerbank, J. S. 1858. On the anatomy and physiol-
ogy of the Spongiadae. Part 1: On the spic-
ules.—Philosophical Transactions of the Royal
Society of London 148:279-—332.
. 1864. A Monograph of the British Spongia-
dae, vol. 1. Ray Society, London, 290 pp.
. 1867. On Alcyoncellum speciosum.—Proceed-
ings of the Zoological Society of London 1867:
351-359.
Gray, J. E. 1867. Notes on the arrangement of sponges,
with the description of some new genera.—Pro-
ceedings of the Zoological Society of London
1867:492-558.
Ijima, I. 1902. Studies on the Hexactinellida. Contri-
bution II. (The genera Corbitella and Hetero-
tella).—Journal of the College of Science, To-
kyo Imperial University 17:1—34.
Kirkpatrick, R. 1910. On Hexactinellida sponge spic-
ules and their names.—Annals and Magazine of
Natural History (8)5:208—213.
Reiswig, H. M., & G. O. Mackie 1983. Studies on
hexactinellid sponges III. The taxonomic status
of Hexactinellida within the Porifera.—Philo-
sophical Transactions of the Royal Society of
London B 301:419—428.
Schmidt, O. 1870. Grundziige einer Spongien-fauna
des Atlantischen Gebietes. Engelmann, Leipzig,
88 pp.
Schulze, E E. 1886. Uber den Bau und das System der
Hexactinelliden.—Abhandlungen der K6nig-
lichen Akademie der Wissenschaften zu Berlin,
Physikalisch-Mathematisch Classe 1886:1—97.
Zittel, K. A. 1877. Studien tiber fossile Spongien; I
Abteilung, Hexactinellidae.—Abhandlungen
der K6niglich Bayerischen Akademie der Wis-
senschaften, Mathematisch-Physikalisch Classe
13:1—63.
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CONTENTS
Taxonomy and evolutionary relationships of Phillips’ small-eared shrew, Cryptotis phillipsii
(Schaldach, 1966), from Oaxaca, Mexico (Mammalia: Insectivora: Soricidae)
Neal Woodman and Robert M. Timm
Two new species of flightless rails (Aves: Rallidae) from the Middle Pleistocene “crane fauna” of Bermuda
Storrs L. Olson and David B. Wingate
A new species of Anthias (Teleostei: Serranidae: Anthiinae) from the Galapagos Islands, with keys to
Anthias and eastern Pacific Anthiinae William D. Anderson, Jr. and Carole C. Baldwin
Description of two new blenniid fish species: Entomacrodus lemuria from the western Indian Ocean
and E. williamsi from the western Pacific Ocean Victor G. Springer and Ronald A. Fricke
Ammodytoides leptus, a new species of sand lance (Teleostei: Ammodytidae) from Pitcairn Island
Bruce B. Collette and John E. Randall
Neostrengeria binderi, a new species of pseudothelphusid crab from the eastern Andes of Colombia
(Crustacea: Decapoda: Brachyura) Martha R. Campos
Freshwater crabs (Brachyura: Potamoidea: Potamonautidae) from the rainforests of the Central African
Republic, central Africa Neil Cumberlidge and Christopher B. Boyko
The freshwater crabs of the Barbilla National Park, Costa Rica (Crustacea: Brachyura: Pseudothelphus-
idae), with notes on the evolution of structures for spermatophore retention
Gilberto Rodriguez and Ingemar Hedstr6m
Acanthilia, a new genus of leucosioid crabs (Crustacea: Brachyura) from the Atlantic coast of the
Americas Bella S. Galil
Cambarus (Cambarus) davidi, a new species of crayfish (Decapoda: Cambaridae) from North Carolina
John E. Cooper
Two new species of Hyalella (Crustacea: Amphipoda: Hyalellidae) from Death Valley National Park,
California, U.S.A. Adam J. Baldinger, William D. Shepard, and Doug L. Threloff
Caecidotea cumberlandensis, a new species of troglobitic isopod from Virginia, with new records of
other subterranean Caecidotea (Crustacea: Isopoda: Asellidae) Julian J. Lewis
Euphilomedes cooki, a new species of myodocopid ostracode from Moreton Bay, SE Queensland,
Australia Elizabeth Harrison-Nelson and Louis S. Kornicker
A new record of Cornechiniscus madagascariensis Maucci, 1993 (Tardigrada: Echiniscidae) from
India Wataru Abe and Masatsune Takeda
Ophryotrocha lipscombae, a new species and a possible connection between ctenognath and labidognath-
prionognath eunicean worms (Polychaeta) Hua Lu and Kristian Fauchald
Aphrodita bisetosa (Polychaeta: Aphroditidae), a new species of sea mouse from the southeastern Pacific
Ocean off central Chile Nicolas Rozbaczylo and Elba Canahuire
Exogone breviantennata Hartmann-Schréder, 1959 (characters emended) (Annelida: Polychaeta: Syllidae),
a new record for the Bahamas with a key to selected Exogone species
Robert Zottoli and Charlene D. Long
Podarke aberrans Webster & Benedict, 1887 - resolution, with descriptions of two new species in the
genus Microphthalmus (Annelida: Polychaeta) Nathan W. Riser
Parapionosyllis cabezali, a new species of Exogoninae (Polychaeta: Syllidae) from Spain
Julio Parapar, Guillermo San Martin, and Juan Moreira
Paralarval gonatid squids (Cephalopoda: Oegopsida) from the Mid-North Atlantic Ocean
Luisa I. Falcon, Michael Vecchione, and Clyde F. E. Roper
Parabuccinum, a new genus of Magellanic buccinulid (Gastropoda: Neogastropoda), with a description
of a new species M. G. Harasewych, Yuri I. Kantor, and Katrin Linse
Porites arnaudi, a new species of stony coral (Anthozoa: Scleractinia: Poritidae) from oceanic islands
of the eastern Pacific Ocean Héctor Reyes-Bonilla and Juan P. Carricart-Ganivet
A new species of Heterotella (Porifera: Hexactinellida: Euplectellidae) from the West Indies
Henry M. Reiswig
Front cover—from this issue, p. 375.
339
356
369
386
397
401
406
420
426
431
443
458
464
480
486
493
500
514
526
552
542
561
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